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In Trinidad, West Indies the social wasps Angiopolybia pallens and Polybia rejecta come to foam nests of the frog Engystomops pustulosus, where they prey on eggs and tadpoles. Frog offspring in early-stage foam nests appear to be almost immune to predation by the wasps, but they become more vulnerable as the nests age and lose definition.

Among amphibians, conspecific chemical communication has been widely studied in Caudata. Adult anurans, by contrast, have received less attention. Recently, it was shown that chemical scents are also relevant for adult anuran intraspecific communication. In this context, we evaluate whether females of the four-eyed frog (Pleurodema thaul) respond to conspecific male scents. We carried out a double choice experiment in a Y-maze. Females were repeatedly presented with the scents of several males versus distilled water. To extract the scent from males, we acoustically stimulated males and then used the water from their aquaria for the experiments. Our data suggest that females are capable of responding behaviourally to male scents, since they spent longer periods in the zones with male scent, rather than in zones with water. We propose that under natural breeding conditions, females of P. thaul may use either their chemical sense or chemical cues to facilitate their encounters with males.

Made available in DSpace on 2020-12-12T02:22:06Z (GMT). No. of bitstreams: 0 Previous issue date: 2020-01-01 The Neotropical frog genus Pseudopaludicola includes 25 species distributed throughout South America. Herein we review the taxonomic status of P. parnaiba relative to P. canga and the specific identity of the population treated in previous studies as Pseudopaludicola sp. 3 from Barreirinhas in the Brazilian state of Maranhão. The lack of differentiation in advertisement call, morphology, and mitochondrial markers from topotypes and different populations rejects the status of P. parnaiba and Pseudopaludicola sp. 3 from Barreirinhas as distinct species. For these reasons, we suggest to formally consider P. parnaiba as a junior synonym of P. canga. We also found that a population previously reported as P. facureae from central Brazil (Palmeiras de Goiás, Goiás) corresponds to a cryptic species that we describe here as a new species. Lastly, we provide for the first time the phylogenetic positions of P. giarettai, P. llanera and P. pusilla. Laboratório de História Natural de Anfíbios Brasileiros (LaHNAB) Departamento de Biologia Animal Instituto de Biologia Universidade Estadual de Campinas (UNICAMP) Laboratório de Taxonomia e Sistemática de Anuros Neotropicais (LTSAN) Instituto de Ciências Exatas e Naturais do Pontal (ICENP) Universidade Federal de Uberlândia (UFU) Instituto de Biologia Universidade Estadual de Campinas (UNICAMP) Laboratório de Herpetologia Departamento de Biodiversidade e Centro de Aquicultura (CAUNESP) Instituto de Biociências Universidade Estadual Paulista (UNESP) Laboratório de Herpetologia Departamento de Biodiversidade e Centro de Aquicultura (CAUNESP) Instituto de Biociências Universidade Estadual Paulista (UNESP)

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"]}

Physalaemus claptoni, new species urn:lsid:zoobank.org:act: 82FFC419-8465-4403-ADE9-289FDED82D92 Holotype (Figs. 1–3; Table 1). UFMG 16784 adult male, November 15, 2014, Serra do Cipó, Fazenda do Chiquito (19.166917° S, 43.529000° W; 1321 m above sea level; datum WGS84), Municipality of Santana do Riacho, State of Minas Gerais, Brazil, F. Leal and H. Thomassen. Paratypes (Figs. 4–6, Table 1). Eighteen adult males: UFMG 16773, 16774, 16776–16783, 16785–16792; one adult female: UFMG 16775, all collected with the holotype by F. Leal and H. Thomassen. Six adult males: UFMG 5418–5423, collected at the type locality on November 1, 2010 by P. C. A. Garcia and F. Natali. One adult male: UFMG 5585, collected at the type locality on November 3, 2010 by F. Leal and P. Taucce. Referred specimens. Two adult males: UFMG 12999–13000, collected at the type locality on December 1, 2011 by P. C. A. Garcia; the individuals were subjected to cytogenetic analyses. One adult male: UFMG 10120, collected at the type locality on January 28, 2012 by P. C. A. Garcia, J. Thompsom, and L. Durães. Diagnosis. The new species is distinguished from all the congener species by the following combination of character states: (1) presence of an arrow-shaped blotch on the dorsum of the body (Figs. 1A, 3, 4, 6A); (2) presence of a median longitudinal light stripe over the urostyle region (Figs. 1A, 3, 4, 6A); (3) belly and ventral surface of thighs marbled with dark gray irregularly shaped blotches, on a bluish background (Fig. 5); (4) absence of reddish coloration over the axillar and inguinal regions in live individuals; (5) bright orange dots scattered on the dorsum of the head, upper lip, and body, in live individuals (Figs. 3, 4); (6) ventral surface of hand and foot red in live individuals; (7) small size (adult male SVL= 16.2–18.2 mm, Table 1); (8) presence of a brown, non-divided, nuptial pad in males (Fig. 7A); (9) END/ED in males ranging from 0.85–0.93 (Table 1); (10) supernumerary tubercles on foot absent (Fig. 2D); (11) tarsal fold absent (Fig. 1B); (12) tarsal tubercle absent; (13); texture of posterior region of belly and ventral surface of thigh smooth (Fig. 1B); (14) advertisement call composed of two note types (note A + B); (15) advertisement call duration of 0.80– 1.28 s (Fig. 8A; Table 2); (16) note A with ascending amplitude until mid-note then descending towards the end of the note; (17) note B with pulses arranged into 5–7 groups; (18) dominant frequency of note A from 1734.4–2765.6 Hz; and (19) dominant frequency of note B from 1507.3–2859.4 Hz (Fig. 8). Comparison with other species. The presence of an arrow-shaped blotch on the dorsum of the body distinguishes Physalaemus claptoni sp. nov. from all the species of the Physalaemus cuvieri Clade (sensu Lourenço et al. 2015). Additionally, the absence of tarsal tubercle distinguishes P. claptoni sp. nov. from the species of the Physalaemus cuvieri Clade, except P. lateristriga, P. olfersii, P. orophilus, and P. maximus of the Physalaemus olfersii species group, from which the new species is promptly distinguishable by the smaller size of adult males at 16.2–18.2 mm (P. lateristriga, P. olfersii, P. orophilus, and P. maximus combined SVL of adult males= 23.1–44.8 mm). Within the P. signifer Clade, the presence of a longitudinal light stripe over the urostyle region distinguishes P. claptoni sp. nov. from all the other species, except from P. rupestris (Fig. 9, S 1), which also has this stripe over the urostyle region (Fig. 9A). By having belly and ventral surface of thigh marbled with dark gray irregularly shaped blotches on a bluish background in live specimens, P. claptoni sp. nov. is set apart from P. angrensis, P. atlanticus, P. nanus, and P. spiniger (belly with flashy orange blotches in those species), from P. maculiventris (which has a pale chest and belly, with posterior region of belly and ventral surface of thigh showing bold black blotches), and from P. obtectus (which has belly with flashy red blotches). The absence of reddish coloration over the axillar and inguinal regions in live individuals distinguishes P. claptoni sp. nov. from P. deimaticus and P. erythros (reddish coloration present over the axillary and inguinal regions on those species). The bright orange dots scattered on the dorsum of the head, upper lip, and body in live specimens of P. claptoni sp. nov. distinguishes it from all the other species within the P. signifer Clade. Ventral surfaces of hand and foot are red in live individuals, which distinguishes P. claptoni sp. nov. from P. angrensis, P. atlanticus, P. maculiventris, P. spiniger (ventral surface of hand and foot orange in these species), and from P. bokermanni, P. caete, P. camacan, P. irroratus, P. moreirae, P. nanus, and P. nattereri (which lack aposematic coloration on ventral surfaces of hand and foot). The small size of P. claptoni sp. nov. adult males at 16.2–18.2 mm promptly distinguishes it from P. atlanticus, P. caete, P. camacan, P. crombiei, P. deimaticus, P. erythros, P. irroratus, P. maculiventris, P. moreirae, P. nattereri, and P. obtectus (combined SVL= 18.9–50.6 mm). The presence of a non-divided nuptial pad in males distinguishes P. claptoni sp. nov. from P. angrensis, P. atlanticus, P. bokermanni, P. caete, P. camacan, P. crombiei, P. deimaticus, P. erythros, P. moreirae, P. nanus, P. obtectus, P. rupestris (Fig. 7B), P. signifer, and P. spiniger (nuptial pad divided in those species). Additionally, P. claptoni sp. nov. is distinguished from P. rupestris by having brown nuptial pad (nuptial pad white-cream in P. rupestris; Fig. 7B) and by the proportion between END/ED in males ranging from 0.85–0.93 (END/ED of P. rupestris males ranges from 0.75–0.82). The absence of supernumerary tubercles on foot distinguishes P. claptoni sp. nov. from P. angrensis, P. caete, P. camacan, P. crombiei, P. irroratus, P. moreirae, P. signifer, and P. spiniger (supernumerary tubercles present on foot of those species). The absence of a tarsal fold distinguishes P. claptoni sp. nov. from P. angrensis, P. atlanticus, P. bokermanni, P. caete, P. camacan, P. crombiei, P. irroratus, P. moreirae, P. nanus, P. obtectus, P. signifer, and P. spiniger (tarsal fold present in those species). The smooth texture of posterior region of belly and ventral surfaces of thigh in P. claptoni sp. nov. distinguishes it from P. camacan and P. irroratus (posterior region of belly and ventral surface of thigh granulated in those species). The advertisement call of P. claptoni sp. nov. is composed of two note types (notes A and B), setting it apart from P. atlanticus, P. angrensis, P. bokermanni, P. camacan, P. crombiei, P. erythros, P. maculiventris, P. moreirae, P. obtectus, and P. signifer (advertisement call has only one note type in those species). The advertisement call of P. claptoni sp. nov. and P. rupestris are both composed of two note types (A and B), however are promptly distinguishable from each other by its duration, which ranges from 0.80– 1.28 s in P. claptoni sp. nov. and from 1.51– 2.20 s in P. rupestris. Additionally, note A of P. claptoni sp. nov. has pulses with an ascending amplitude until mid-note and then descends towards the end of the note, while note A of P. rupestris has only descending amplitude modulation; note B of P. claptoni sp. nov. has pulses arranged in 5 to 7 pulse groups, whereas note B of P. rupestris has pulse grouping only at the onset and final portion of the note. Dominant frequency of note A in P. claptoni sp. nov. varies from 1734.4–2765.6 Hz and in P. rupestris from 3046.9–3421.9 Hz. Dominant frequency of note B in P. claptoni sp. nov. ranges from 1507.3–2859.4 Hz and in P. rupestris from 3046.9–3421.9 Hz. Description of the holotype. Adult male, SVL 17.3 mm (Fig. 1–3). Head slightly longer than wide. Head width (32.4% SVL) slightly shorter than head length (34.7% SVL). Snout short, rounded in dorsal and lateral views (Fig. 2). Canthus rostralis distinct, rounded; loreal region slightly concave. Snout protruding beyond lower jaw. Nostril dorsolaterally oriented, faintly protruding. Internarial region slightly concave; top of the head flat. Eye slightly prominent, antero-laterally oriented, its diameter 11% bigger than END. Tympanum indistinct externally. Supratympanic fold distinct, thick, extending from the posterior corner of the eyes to the shoulders. Dentigerous process of vomer absent. Premaxillary and maxillary teeth absent. Choanae rounded, separated from each other by a distance as large as four times its diameter. Tongue elliptical, elongated, slightly constricted in its middle, free around lateral and posterior margin. Vocal slit present, longitudinal, originating on the sides of the tongue in its anterior third, and extending towards the corner of the mouth. Vocal sac single and subgular, weakly differentiated externally, slightly folded over the chest (in preservative). Dorsolateral fold present, weakly distinct, from the posterior corner of the eye to the inguinal region. Forearm very hypertrophied in relation to upper arms; upper arms slender, short. Fingers thick, without webs, relative lengths I=II=III Measurements (in mm) of the holotype: SVL 17.3; HL 6.0; HW 5.6; ED 1.6; AMD 3.1; ESD 2.5; END 1.4; IND 1.4; THL 7.6; TL 7.0; FL 11.0; HAL 4.1. Color of the holotype in life. Dorsal background color of the head, body, upper arms, and forearms light orange. Interorbital blotch leaden gray colored, irregularly shaped, curved towards the body, extending from the superior edge of one eyelid to the other; anterior margin well-defined, bordered by a very thin white line; posterior margin diffuse. Dorsum of the body with an arrow-shaped blotch medially located, gray leaden, and outlined by a very thin white line; its posterior portion extends transversely toward the inguinal gland, covering the sacrum region; a small white spot is displayed on the middle of the arrow-head and there is a longitudinal light stripe over the urostyle region. Black ocelli cover 75% of the inguinal gland on its posterior portion. The dorsal and hidden surfaces of thigh, tibia and tarsus leaden gray colored; dorsal surface of the toes is light orange; transversal black bar on the dorsal surfaces of thigh, tibia, and foot, aligned to each other, slightly invading the posterior surface of the thigh and ventral surface of the tibia. Irregularly shaped black blotches on the heel, knee, and ventral surface of tarsus. Longitudinal elongated black blotch on the posterior surface of upper arm, extending slightly dorsally to the elbow. Hand and distal half of the forearm with small black spots and scattered orange dots. Dorsolateral gray leaden-colored blotch, extending from the posterior margin of the eye, passing over the supratympanic fold and almost reaching the inguinal region; stained by black spots on the supratympanic fold and on the posterior half of the flank. Canthus rostralis, upper half of the loreal region and lateral surfaces of the snout stained by small black spots; supralabial region bright orange colored. Black blotches under the eye separated from each other by an orange surface. Dorsum of the body and head with scattered bright orange dots. Ventral surfaces of the limbs marbled with dark gray and bluish irregular shaped spots; black blotch over the glandular tissue on the ventrolateral edge of forearm. Black bar above the cloaca, curved down toward the posterior surface of the thighs. Gular region and chest dark gray, stained with scattered bluish dots. Belly and ventral surface of thigh marbled with dark gray irregularly shaped blotches, on a bluish background. Ventral surface of the hand red; toes I and II, external metatarsal tubercles, internal metatarsal tubercle and subarticular tubercles of the foot red; plantar region and region between subarticular tubercles of the toes III, IV and V black with scattered irregular shaped whitish spots. Iris orange, adorned with black vermiculation; pupil black and horizontal. Color of the holotype in alcohol 70%. In preservative, all the dorsal coloration took on dark gray shades; the dorsal blotches become barely distinguishable; all the orange coloration faded. Ventrally, the black and dark gray colorations become brownish dark gray, bluish coloration become pale white; and the red coloration of the hand and foot become pale white. Variation. Measurements of 26 adult males and one adult female are presented in Table 1. The adult female (Fig. 6) differs from adult males by having slender forearm; vocal slit absent; nuptial pad absent; and HL and HW smaller in relation to SVL. The left hand of the female paratype is malformed, with smaller fingers. In life, color of dorsal background of head, body, upper arm, and forearm may vary from pale yellow to bright orange; color of interorbital, dorsal and dorsolateral blotches as well as dorsal and hidden surfaces of thigh, tibia and tarsus may be brown or dark gray; the orange dots on the dorsum of head and body vary in number and intensity of color, being pinkish in some individuals; the area with orange coloration over upper lips and canthus rostralis varies in size (Fig. 4). Interorbital blotch may be M shaped; tip of the arrow-shaped blotch may be in contact with the interorbital blotch. Median longitudinal light stripe over the urostyle region varies in size and thickness. Irregularly shaped blotches of belly and ventral surface of thigh may vary in size and number. Iris varies from pale yellow to orange. In preservative, the current (2019) general coloration of specimens collected in 2010 is brownish, whereas the ones collected on 2014 remain grayish. Tympanum may be slightly discernible externally in some individuals. Head may be longer than wide or as long as wide. Dark superficial keratinized layer of nuptial pad may be peeled, but nuptial pad remains visible. Etymology. The specific epithet is used as a noun and honors Eric Clapton, the English rock and blues guitarist, singer, and songwriter. Clapton is widely recognized as one of the best guitarists of all time. Over his career, Clapton’s work has made an outstanding contribution to music, influencing many musicians all around the world. Advertisement call. The advertisement call of Physalaemus claptoni sp. nov. (Fig. 8A, B; Table 2; n=45 calls from five males; Audio S1) consists of two types of multipulsed notes (herein referred as notes A and B) released in sequence. It is formed by 65–93 pulses, has duration ranging from 0.80– 1.28 s (n=45 advertisement calls), interval between notes A and B from 0.10– 0.19 s (n=45 intervals) and between advertisement calls varying from 0.14– 1.38 s (n=41 intervals). Note rate within advertisement call varied from 1.56–2.51 notes/s (n=90 notes). Note A (n=154 notes) is shorter than B, with duration varying from 0.11– 0.16 s, is formed by 12–17 pulses emitted at a rate of 86.7–119.3 pulses/s. Note A was released at a rate of 0.53–0.63 notes/s. Pulses of note A are homogeneously distributed and have ascending amplitude until mid-note then descending towards the end of the note, being fairly symmetric (peak amplitude from 0.44–0.59). Notes A were also observed being released in sequences without note B, and, sometimes, between advertisement calls. In this case, we considered each note A as a single call. The interval between single A note calls emitted in sequences ranged from 0.7– 1.9 s (n=113 intervals). Note B (n=45 notes, from four males) duration varies from 0.67– 1.01 s and is formed by 53–81 pulses emitted at a rate of 67.5–92.1 pulses/s. Note B was released at a rate of 0.32–0.61 notes/s. Pulses of note B are arranged in 5–7 pulses group, with relative amplitude lower than that of note A. Within those groups, pulses are also homogeneously distributed and with ascending amplitude until its midpoint, then descending towards the end of the pulse group. The first and second pulse groups are generally closer and even in contact to each other than are the others (See Fig. 8 A). The other pulse groups of note B may have a complete amplitude modulation in between groups, or not. In the latter case, there are a few isolated and low-amplitude pulses in between groups. Frequency bandwidth of note A (n=154 notes) is between 1453.1–3140.6 Hz and of note B (n=45 notes) is between 1335.1–3375.0 Hz. Dominant frequency of note A (n=154 notes) ranged from 1734.4–2765.6 Hz and of note B (n=45 notes) from 1507.3–2859.4 Hz. Phylogenetic inferences and genetic distances. The phylogenetic analysis recovered one most parsimonious tree, with 5853 steps. In the inferred cladogram, Physalaemus claptoni sp. nov. was nested inside the P. deimaticus species group, a clade that was highly supported (bootstrap value of 100%; Fig. 10). In this analysis, the P. signifer Clade and the P. cuvieri Clade were recovered

We describe a new species of Physalaemus assigned to the Physalaemus signifer Clade, and it is morphologically similar to P. rupestris, from the highlands of the Serra do Cipó in the southern Espinhaço Range, State of Minas Gerais, Southeastern Brazil. The new species is diagnosed by using the following combination of character states: presence of an arrow-shaped blotch on the dorsum of body; presence of a median longitudinal light stripe over urostyle region; belly and ventral surface of thigh marbled with dark gray irregularly shaped blotches on a bluish background; absence of reddish coloration over axillary and inguinal regions in live individuals; bright orange dots scattered over head, upper lip, and dorsum of body in live individuals; ventral surface of hand and foot red in live individuals; small size (adult male SVL=16.2–18.2 mm); presence of brown, not divided, nuptial pad in males; END/ED in males ranging from 0.85–0.93; supernumerary tubercles on foot absent; tarsal fold absent; tarsal tubercle absent; texture of posterior region of belly and ventral surface of thigh smooth; advertisement call composed of two note types (note A + B); advertisement call duration of 0.80–1.28 s; note A with ascending amplitude until mid-note then descending towards the end of the note; note B with pulses arranged in 5–7 groups; and dominant frequency of note A from 1734.4–2765.6 Hz and of note B from 1507.3–2859.4 Hz. A phylogenetic analysis based on mitochondrial DNA sequences recovered the new species nested within the Physalaemus deimaticus species group. Additionally, we redescribe the call of Physalaemus rupestris and provide a review of the geographic distribution and conservation status of the species belonging to the P. deimaticus species group.

Physalaemus deimaticus species group distribution and conservation Physalaemus deimaticus was described in 1986 based on three individuals collected from the highlands of Serra do Cip��, with two of them collected in 1972 and one collected in 1982 (Sazima & Caramaschi 1986). The species was missing until 2010, when two individuals were collected at the type locality by M.T. Rodrigues. Likewise, P. erythros was described in 2003 on the basis of three individuals that were collected from the Parque Estadual do Itacolomi (Caramaschi et al. 2003). Ba��ta & Silva (2009) extended its distribution based on a single individual collected at the Reserva Particular do Patrim��nio Natural Santu��rio do Cara��a (RPPN Cara��a), in the Municipality of Catas Altas, ca. 33 km northward from the type locality. The apparently low abundance and poor knowledge about these species in terms of biology and distribution led both of them to be categorized as Deficient Data (DD) in the Brazilian Red List of Threatened Amphibian Species (ICMBio 2014). The National Action Plan for Threatened Extinction Reptiles and Amphibians Conservation of Espinha��o Range (PAN Espinha��o) (MMA 2012, 2016) recommended the inventory of areas of potential occurrence of these DD species. Encouraged by this, we have searched for new populations of Physalaemus. deimaticus and P. erythros over the last few years (see Appendix I). For P. deimaticus we discovered two new populations: one in the Municipality of Diamantina, 112km northward from its type locality, representing its northern limit of distribution; the others were in the Municipalities of Serro and Santo Ant��nio do Itamb�� in the Parque Estadual Pico do Itamb��, ca. 98 km northward from its type locality, representing its eastern limit of distribution. We also registered P. deimaticus at its type locality in six different streams. All these P. deimaticus populations were relatively abundant, could be well sampled, and were included in our phylogenetic analysis. For Physalaemus erythros, we also discovered two new populations: one in the Municipality of Bar��o de Cocais, 59 km northward from its type locality and 27 km northward from the RPPN Cara��a, representing its northern limit of distribution; and the other in the Municipality of Santa B��rbara, at the Parque Nacional do Gandarela, 36 km northward from its type locality and 20 km west of the RPPN Cara��a, representing its western limit of distribution. We also registered P. erythros at its type locality in four different streams. All the registered populations of P. erythros were relatively abundant and could be sampled well and were included in our phylogenetic analysis, except for the one from Parque Nacional do Gandarela, from which we collected only one female from a pitfall trap. In view of the data we provide here on the distribution of these species, with Physalaemus deimaticus occurring in two protected areas and P. erythros in three protected areas, there is no evidence of threats that could lead to their extinction. Therefore, we suggest that they should be considered as least concern (LC) species. Physalaemus claptoni sp. nov. occurs geographically close (ca. 10 km north) to the type locality of P. deimaticus in the Espinha��o Range (Fig. 11). However, they reproduce in distinct habitats. While P. claptoni sp. nov. reproduces in small still water collections associated with forest edges, P. deimaticus uses small temporary rock streams surrounded by grasslands. Physalaemus rupestris is known from the mountains of Serra do Ibitipoca and Serra Negra in the Mantiqueira Range (Caramaschi et al. 1991; Oliveira et al. 2009), and it is the only species of the P. deimaticus group distributed outside the Espinha��o Range. Despite occurring in a distinct mountain range (Mantiqueira Range) where rock formations are more commonly made up of gneiss and granite (Vasconcelos 2011), P. rupestris is only found in some of the few quartzitic mountains of this range that harbor the same vegetation found in the Espinha��o Range, namely, campo rupestre (Vasconcelos 2011; Leal, F. personal observation). The occurrence of closely related species that are allopatrically distributed in the Mantiqueira (P. rupestris) and Espinha��o (P. claptoni sp. nov.) ranges suggests a historic connection between these highlands of eastern Brazil, a hypothesis that has already been pointed out (Maxson & Heyer 1982; Cruz & Feio 2007) and is also corroborated by the existence of species with distributions restricted to the Espinha��o and Mantiqueira ranges (Pirani et al. 2010; Taucce et al. 2012). Further phylogeographical studies based on larger samples of specimens and genes (including nuclear markers) could help us to elucidate the historical and evolutionary processes underlying the occupation of these two mountain ranges by the P. deimaticus species group. The new species has not been registered within strictly protected reserves, although it occurs between two of them, at 7 km from the Parque Nacional da Serra do Cip�� (33.8 ha) and 5 km from the Parque Estadual Serra do Intendente (13.5 ha). The type locality is located in a private area where wood extraction has affected the structure of the forest fragment for years, although lately this activity has declined. Moreover, fire and cattle breeding have been disturbance agents for decades within the entire Serra do Cip��, including the type locality of Physalaemus claptoni sp. nov. Even though several individuals of P. claptoni sp. nov. were found during our fieldwork from 2010 to 2017, we could not evaluate the possible effects of these disturbance activities on this species in a medium-long term. Since the compilation of the anuran endemic species from the Espinha��o Range made by Carvalho et al. (2013), who listed 37 taxa, four new species with distribution restricted to the mountain range have been described: Crossodactylodes itambe Barata, Santos, Leite, & Garcia, 2013, Sphaenorhynchus canga Araujo-Vieira, Lacerda, Pezzuti, Leite, Assis, & Cruz, 2015, Scinax montivagus Junc��, Napoli, Nunes, Merc��s, and Abreu, 2015, Odontophrynus juquinha Rocha, Sena, Pezzuti, Leite, Svartman, Rosset, Baldo, and Garcia, 2017, and Physalaemus claptoni sp. nov. (present study). On the other hand, Ololygon tripui Louren��o, Nascimento & Pires, 2009 and Bokermannohyla nanuzae (Bokermann & Sazima, 1973b) lost their endemic status because they were found outside the Espinha��o (Silva et al. 2013) or were synonymized with species that occur elsewhere (Walker et al. 2015), respectively. Therefore, 39 anuran species should be considered endemic to the Espinha��o Range at present. Acknowledgments We thank all those who provided assistance in the field expeditions, especially Hans Thomassen who helped collecting most of type series of the new species described and topotypes of P. deimaticus, Renato N. Feio who helped collecting and recording many of the P. rupestris analyzed, and Pedro C. Rocha who helped collecting most of the P. deimaticus, P. erythros, and many of the P. rupestris analyzed. We also thank Pedro C. Rocha for making the distribution map (Fig. 11). We thank Igor R. Fernandes for help with the analysis of nuptial pads. We thank Miguel R. Trefaut for providing information regarding the specimens of P. deimaticus collected by him. We thank the collection curators and lab staff of UFV (Renato N. Feio, Carla S. Guimar��es), MNRJ (Jos�� P. Pombal Jr., Manoela W. Cardoso), and CFBH (C��lio F. B. Haddad, Nadya Pupin). Scientific collecting permits were issued by the Brazilian Instituto Chico Mendes de Conserva����o da Biodiversidade (SISBIO licenses 42369-1, 51066-2, 22551-1, 22551-2, 35826-1) and Instituto Estadual de Florestas (IEF UC: 146/12, 147/12). Felipe S. F. Leite thanks to Funda����o de Amparo �� Pesquisa do Estado de Minas Gerais (FAPEMIG, Process APQ-2067-14; APQ-01796-15; RDP-00004-17). Paulo C. A. Garcia thanks the productivity grant provided by the Conselho Nacional de Desenvolvimento Cient��fico e Tecnol��gico (CNPq), Funda����o de Amparo a Pesquisa do Estado de Minas Gerais (FAPEMIG, Process APQ- 03462-09; RDP-00053-10; PPM-00515-10). Luciana B. Nascimento thanks Conselho Nacional de Desenvolvimento e Pesquisa (CNPq, Process 479457/2012-03), Funda����o de Amparo �� Pesquisa do Estado de Minas Gerais (FAPEMIG, Process APQ-2067-14), and Fundo de Incentivo �� Pesquisa - PUC Minas (FIP/PUC Minas). Luciana B. Louren��o thanks Funda����o de Amparo �� Pesquisa do Estado de S��o Paulo (FAPESP, Process #2011/09239-0)., Published as part of Leal, Fernando, Leite, Felipe S. F., Da Costa, William P., Nascimento, Luciana B., Louren��o, Luciana B. & Garcia, Paulo C. A., 2020, Amphibians from Serra do Cip��, Minas Gerais, Brasil. VI: A New Species of the Physalemus deimaticus Group (Anura, Leptodactylidae), pp. 306-330 in Zootaxa 4766 (2) on pages 323-325, DOI: 10.11646/zootaxa.4766.2.3, http://zenodo.org/record/3764071, {"references":["Caramaschi, U., Feio, R. N. & Guimaraes Neto, A. S. (2003) A new brightly colored species of Physalaemus (Anura: Leptodactylidae) from Minas Gerais, southeastern Brazil. Herpetologica, 59, 519 - 524. https: // doi. org / 10.1655 / 02 - 102","Baeta, D. & Silva, D. H. (2009) Amphibia, Anura, Leiuperidae, Physalaemus erythros Caramaschi, Feio and Guimaraes-Neto, 2003: Distribution extension. Check List, 5, 812 - 814. https: // doi. org / 10.15560 / 5.4.812","Caramaschi, U., Carcerelli, L. C. & Feio, R. N. (1991) A new species of Physalameus (Anura: Leptodactylidae) from Minas Gerais, southeastern Brazil. Herpetologica, 47, 148 - 151.","Oliveira, E. F., Tolledo, J. & Feio, R. N. (2009) Amphibia, Anura, Physalaemus rupestris Caramaschi, Carcerelli and Feio, 1991: Distribution extension and geographic distribution map. Check List, 5, 815 - 818. https: // doi. org / 10.15560 / 5.4.815","Vasconcelos, F. V. (2011) O que sao campos rupestres e campos de altitude nos topos de montanha do Leste do Brasil? Revista Brasileira de Botanica, 34, 241 - 246. https: // doi. org / 10.1590 / S 0100 - 84042011000200012","Maxson, L. R. & Heyer, W. R. (1982) Leptodactylid frogs and the Brasilian Shield: an old and continuing adaptive relationship. Biotropica, 14, 10 - 15. https: // doi. org / 10.2307 / 2387754","Cruz, C. A. G. & Feio, R. N. (2007) Endemismos em anfibios em areas de altitude na Mata Atlantica no sudeste do Brasil. In: Nascimento, L. B. & Oliveira, M. E. (Eds.), Herpetologia no Brasil II. Sociedade Brasileira de Herpetologia, Sao Paulo, pp. 117 - 126.","Pirani, R. M., Mangia, S., Santana, D. J., Assis, B. & Feio, R. N. (2010) Rediscovery, distribution extension and natural history notes of Hylodes babax (Anura, Hylodidae) with comments on Southeastern Brazil Biogeography. South American Journal of Herpetology, 5, 83 - 88. https: // doi. org / 10.2994 / 057.005.0202","Taucce, P. P., Leite, F. S., Santos, P. S., Feio, R. N. & Garcia, P. C. (2012) The advertisement call, color patterns and distribution of Ischnocnema izecksohni (Caramaschi and Kisteumacher, 1989) (Anura, Brachycephalidae). Papeis Avulsos de Zoologia, 52, 112 - 120. https: // doi. org / 10.1590 / S 0031 - 10492012000900001","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","Junca, F. A., Napoli, M. F., Nunes, I., Merces, E. A. & Abreu, R. O. (2015) A New Species of the Scinax ruber Clade (Anura, Hylidae) from the Espinhaco Range, Northeastern Brazil. Herpetologica, 71, 299 - 309. https: // doi. org / 10.1655 / HERPETOLOGICA-D- 14 - 00032","Rocha, P. C., Sena, L. M. F., Pezzuti, T. L., Leite, F. S. F., Svartman, M., Rosset, S. D, Baldo, D. & Garcia, P. C. A. (2017) A new diploid species belonging to the Odontophrynus americanus species group (Anura: Odontophrynidae) from the Espinhaco range, Brazil. Zootaxa, 4329 (4), 327 - 350. https: // doi. org / 10.11646 / zootaxa. 4329.4.2","Lourenco, A. C. C, Nascimento, L. B. & Pires, M. R. S. (2009) A new species of the Scinax catharinae species group (Anura: Hylidae) from Minas Gerais, southeastern Brazil. Herpetologica, 65, 468 - 479. https: // doi. org / 10.1655 / 07 - 088.1","Bokermann, W. C. A. & Sazima, I. (1973 b) Anfibios da Serra do Cipo, Minas Gerais, Brasil. II: Duas especies novas de Hyla (Anura, Hylidae). Revista Brasileira de Biologia, 33, 521 - 528.","Silva, G. R., de Luna-Dias, C., Hepp, F. S. F. S. & Carvalho-e-Silva, S. P. (2013) First record of Scinax tripui Lourenco, Nascimento and Pires, 2010 (Amphibia: Anura: Hylidae) from Espirito Santo state, Brazil. Check List, 9, 645 - 646. https: // doi. org / 10.15560 / 9.3.645","Walker, M., Lourenco, A. C. C., Pimenta, B. V. S. & Nascimento, L. B. (2015) Morphological variation, advertisement call, and tadpoles of Bokermannohyla nanuzae (Bokermann, 1973), and taxonomic status of B. feioi (Napoli & Caramaschi, 2004) (Anura, Hylidae, Cophomantini). Zootaxa, 3937 (1), 161 - 178. https: // doi. org / 10.11646 / zootaxa. 3937.1.8"]}

This study summarizes new and historical records of one of the least known anuran species from the Chilean Altiplano, Pleurodema marmoratum (Duméril & Bibron, 1840). The discovery of a new population in the Altiplano of the Región de Antofagasta extends the known distribution of this species in Chile approximately 100 km south.

Physalaemus olfersii (Lichtenstein & Martens, 1856) We found a single call type for the species, referred to as call A. The call is composed of a single harmonic note with a very long duration and a slight PAM (no silence intervals between pulses). It has an irregular and strong PFM throughout the call. The bands have no general FM or have only a slight FM, which is usually upward. Call A (Fig. 44 A–D and 42C). We examined 13 recordings, a total of 35 minutes, with 332 calls from 31 males. Only some of these calls were measured (see Table 2). Call duration varies from 3.530 to 4.837 s. Call rise and fall are very short and similar to each other in duration. The limit between the call rise and sustain is not clear in calls with triangular envelope (Fig. 44A). There is a long sustain. This segment is usually regular and almost flat (slightly decreasing towards end of the segment – Fig. 44C), but some calls have very inclined sustains, where the beginning of the segment has low amplitude and gradually increases towards its end (Fig. 44A). The amplitude peak is at around the end of the first tenth or at the very end of the call duration. The envelope of the call varies from rectangular (Fig. 44C) to triangular (pointed left; Fig. 44A). More than 50 % of the call energy is concentrated in 46 % of the call duration around the amplitude peak. The call can have a slight PAM (silence intervals are absent between peaks). The rate of the PAM is ca. 11 Hz, forming ca. 50 cycles throughout the call. The cycle rise and fall are similar and the amplitude peak is at the middle of the cycle duration. The call has a harmonic series (Fig. 42C). The fundamental frequency is ca. 150 Hz. The first seven harmonics have very low energy or are absent in the audiospectrogram. There are ca. 12 adjacent emphasized harmonics. The wave periods are usually regular and harmonics are clear throughout the call. However, some calls have deterministic chaos regimes in several parts, mainly at their outset (Fig. 44B). The dominant frequency varies from ca. 1570 to 1870 Hz (Fig. 44B). The dominant harmonic varies from the ninth to the 19 th, but it is usually between the ninth and 12 th (Fig. 42C). There is no clear shift in the relative energy between the bands throughout the call. Most of the call energy is between 1100 and 2150 Hz (12 harmonics). Calls have no general FM (Fig. 44D), or have only a slight general FM, usually upward (Fig. 44B), but sometimes downward. There is usually a short downward FM at the end of the call (Fig. 44B). Additionally, there is a strong PFM throughout the call, which is usually independent, but it is directly proportional and synchronic to PAM when it is present (Fig. 44B, D)., Published as part of Hepp, Fábio & Pombal, José P., 2020, Review of bioacoustical traits in the genus Physalaemus Fitzinger, 1826 (Anura: Leptodactylidae: Leiuperinae), pp. 1-106 in Zootaxa 4725 (1) on page 76, DOI: 10.11646/zootaxa.4725.1.1, http://zenodo.org/record/3612996

Physalaemus carrizorum Cardozo & Pereyra, 2018 We found a single call type for the species, referred to as call A. The call is composed of a single harmonic note, with general downward FM, but with an up-downward FM segment in the first fourth of the call. Call A (Fig. 49 A–D and 42H). We examined one recording, a total of four minutes, with 44 calls from three males. Only some of these calls were measured (see Table 2). Call duration varies from 2.360 to 4.118 s. The envelope of the call is variable. In most calls, rise and fall are similar in duration and shape (exponential) and the sustain is long. Some sustains are regular (Fig. 49C) and others are irregular (Fig. 49A), with short and shallow valleys. In some calls, the limits between the call rise, sustain, and call fall are not clear. Usually, the envelope is divided into two parts with different amplitude levels (Fig. 49A). The amplitude peak is usually at the end of the first seven tenths of the call duration. The envelope varies from elliptic or rectangular (Fig. 49C), to triangular (pointed left; Fig. 49A; rarely pointed right). Due to the asymmetry of some triangular envelopes, the shape resembles an arrow. More than 50 % of the call energy is concentrated in 36 % of the call duration around the amplitude peak. There is no PAM in the call. The call has a harmonic series (Fig. 42H). The fundamental frequency is ca. 460 Hz and approximately the first seven harmonics are emphasized. The wave periods are regular and harmonics are clear throughout the call. The dominant frequency varies from ca. 410 to 2630 Hz (Fig. 49B). The dominant harmonic is the first, fifth, or sixth (Fig. 42H). There is a clear shift in relative energy between the bands; the dominant frequency gets higher toward the end of the call, starting at the first harmonic, moving to the fifth, and ending at the sixth (Fig. 42H, 49B). Most of the call energy is between 350 and 3500 Hz (eight to nine harmonics). The call has a general downward FM (Fig. 49B, D). Additionally, calls have an up-downward FM in the first fourth of the call duration, leading to slightly arc-shaped bands in this part of the call (Fig. 49B, D), and a short downward FM at the end (Fig. 49B). The general downward FM and the initial up-downward FM result in S-shaped harmonics when considering the entire call. Some calls have a slight PFM., Published as part of Hepp, Fábio & Pombal, José P., 2020, Review of bioacoustical traits in the genus Physalaemus Fitzinger, 1826 (Anura: Leptodactylidae: Leiuperinae), pp. 1-106 in Zootaxa 4725 (1) on page 80, DOI: 10.11646/zootaxa.4725.1.1, http://zenodo.org/record/3612996, {"references":["Cardozo, D. E. & Pereyra, M. O. (2018) A new species of Physalaemus (Anura, Leptodactylidae) from the Atlantic Forest of Misiones, northeastern Argentina. Zootaxa, 4387 (3), 580 - 590. https: // doi. org / 10.11646 / zootaxa. 4387.3.10"]}

Physalaemus bokermanni Cardoso & Haddad, 1985 We found two different calls, referred to as call A and B. Calls B were common in recordings in which several males were active and calling at the same night. Calls A and B are composed of harmonics and pulses (i.e., pulse-PAM). Call B has two notes while Call A has only one. The first note of call B is similar to that of call A. The second note of call B is much longer than that of call A and has an envelope with a long and gradual rise. Call A (Fig. 18 A–D and 13E). We examined eight recordings, a total of 12 minutes, with ca. 650 calls from nine males. Only some of these calls were measured (see Table 2). Call duration varies from 0.177 to 0.197 s. The call rise is linear or logarithmic-shaped and longer than the fall, which is usually abrupt and logarithmic-shaped; the amplitude peak is at around the end of the first three fourths of the call duration (Fig. 18A). The envelope of the call is elliptic or triangular (pointed left; Fig. 18A, C). More than 50 % of the energy is concentrated in 41 % of the call duration around the amplitude peak. This call has a strong PAM (with silence intervals present between pulses; Fig. 18 A–D). The rate of the PAM is ca. 35 Hz, forming ca. six pulses throughout the call. The pulse rise is abrupt and much shorter than the fall; the amplitude peak is at the beginning of the pulse (Fig. 18C). The first one or two pulses have much lower amplitude than the others (Fig. 18A). Often, the second and the last pulses are the longest (Fig. 18A, B). Silence intervals are present between pulses, ca. fivefold longer than pulse duration. The first interval is usually much shorter than the others (Fig. 18 A–D). The call has a harmonic series (Fig. 13E). The fundamental frequency is ca. 780 Hz and this band can be present with low energy or absent in the audiospectrograms. The short duration of the pulses makes the bands broad with narrow intervals (Fig. 18B, D). Some pulses can have not very clear harmonics, with considerably deterministic chaos due to the irregularity of the wave periods (Fig. 18B, D). There are jumps of the fundamental frequency between pulses in some calls. The dominant frequency varies from ca. 2840 to 3660 Hz (Fig. 18B). The dominant harmonic varies from the second to the 10 th, but it is usually the fourth. There is no clear shift in the relative energy among the bands throughout the call. Most of the energy is concentrated between 2600 and 3900 Hz (ca. three harmonics). Usually, there is no clear general FM throughout the call, however, in some calls the first two pulses have their energy concentrated in lower frequency bands, making the general FM of the call upward (Fig. 18B). Call B (Fig. 18 E–H and 16C). We examined three recordings, a total of six minutes, with 35 calls from three males. Only some of these calls were measured (see Table 2). Call duration varies from 0.947 to 1.868 s and the call has two different notes. Duration of the second note is ca. 1.0 s. The amplitude, temporal and spectral traits of the first note resemble those of call A, although in call B the first note often has more abrupt rise and fall (Fig. 18E). Usually, there is a silence interval between the notes (Fig. 18E, F). However, in some calls, this interval is perceptible only as a decrease in amplitude. The rise of the second note is logarithmic-shaped and shorter than fall, which is gradual, almost linear; the amplitude peak of the note is at the end of the first tenth of the note duration (Fig. 18E). Due to the very short rise and the long and gradual fall, the envelope of the component B is triangular (pointed right; Fig. 18E). More than 50 % of the energy of the compound call is concentrated in ca. 34 % of the duration around the amplitude peak. Both notes have a strong PAM (there are silence intervals present between pulses; Fig. 18 E–H). The rate of the PAM is similar to that of the call A, ca. 30 Hz, yielding 28 pulses throughout the call. The rate of the PAM is less regular in the second note than in the first one. The pulse rise is abrupt and much shorter than the fall; the amplitude peak is at the beginning of the pulse (Fig. 18G). Some pulses can be twofold longer than the others. At the beginning of the second note the ratio between the silence interpulse-interval and pulse duration is similar to that in component A. The interval becomes longer (pulse duration remains the same) towards the end of the call (i.e., pulse-PAM rate decreases), mainly after the first fourth of the second-note duration (Fig. 18E, G). Spectral traits of the second note are similar to those of call A (Fig. 18F, H; see some quantitative differences in Table 2). There is no general FM in component B (Fig. 18F, H).

Physalaemus erythros Caramaschi, Feio & Guimarães, 2003 We found two different calls, referred to as call A and B. B calls were common in recordings in which several males were active and calling close to each other. B calls were commonly observed after overlapping periods of calls A. Call B differs from call A by its longer duration, higher fundamental frequency, presence of pulse-PAM and PFM. Additionally, the envelope of A calls is elliptic whereas that of the B calls is triangular or rectangular. Call A (Fig. 7 A–D and 4D). We examined 21 recordings, a total of 56 minutes, with ca. 4900 calls from 15 males. Only some of these calls were measured (see Table 2). Call duration varies from 0.041 to 0.077 s. The call rise and fall are similar in duration, rendering an elliptic call envelope; the call fall is gradual, whereas the call rise is steeper. The amplitude peak is at around the middle of the call duration (Fig. 7A, C). More than 50 % of the call energy is concentrated in 39 % of the call duration around the amplitude peak. The call has no PAM. The call is composed of harmonics (Fig. 4D). The harmonics are very close to each other and hardly distinguished due to the low fundamental frequency and the lack of the wave periodicity throughout the call. The fundamental frequency is approximately 250 Hz and this band can be present with low energy or absent in the audiospectrograms. The dominant frequency varies from ca. 1020 to 1160 Hz (Fig. 7B, D). The dominant harmonic varies from the second to the tenth, but it is usually the third. There is no clear shift in the relative energy among the bands throughout the call. Most of the call energy is between 300 and 2000 Hz (ca. seven harmonics). There is no clear general FM in the call but there are subtle irregular FM segments throughout the entire call. Call B (Fig. 7 E–J and 6B). We examined eight recordings, a total of 26 minutes, with 93 calls from eight males. Only some of these calls were measured (see Table 2). Call duration varies from 0.200 to 0.875 s. The call rise is abrupt and a little shorter than the fall, which is also very short but gradual. The call has a long sustain. The amplitude is usually regular throughout the call (Fig. 7H). However, in some calls, the amplitude increases gradually toward the end of the call (Fig. 7E, G). The amplitude peak is at the very end of the call duration (Fig. 7E, G, H). Depending on the slope of the sustain and the difference between the amplitude peaks, the envelope of the call can vary from rectangular (Fig. 7H) to triangular (pointed left; Fig. 7E, G). More than 50 % of the call energy is concentrated in 29 % of the call duration around the amplitude peak. The call has a weak to intermediate PAM (there is no silence interval between the peaks; Fig. 7E, G, H). The rate of the PAM is ca. 27 Hz, forming ca. 13 amplitude peaks throughout the call. The envelope of this PAM cycles is variable but the amplitude peak is at the middle of the cycle. The call is composed of harmonics (Fig. 6B). Usually the harmonics are clear, however, eventual decrease in the wave periodicity makes the harmonics less clear with deterministic chaos regimes. Audiospectrograms with relatively broad filter bandwidths (e.g., above 100 Hz) can show wave peaks in some parts of the call with low fundamental frequencies (minimum 107 Hz; see Table 2), as broadband pulses (instantaneously high sound-pressure effect; see Littlejohn 2001). The fundamental frequency is around 250 Hz and it is usually absent in the audiospectrograms. The dominant frequency varies from ca. 840 to 1780 Hz (Fig. 7F, I, J). The dominant harmonic varies from the third to the ninth, but it is usually the third and fourth harmonic. There is no clear shift in the relative energy among the bands throughout the call. Most of the call energy is between 800 and 1600 Hz (three to four harmonics). The frequency bands have a general upward FM throughout the call and a short downward FM at the end (Fig. 7F, I, J). There are irregular PFM segments throughout the entire call; these segments are usually synchronic and directly proportional to the PAM (Fig. 7 E–J).

Physalaemus riograndensis Milstead, 1960 We found a single call type for the species, referred to as call A. The call is composed of a single harmonic note with high fundamental frequency (ca. 1000 Hz). It has a general downward FM throughout the call, with an up-downward FM segment in the its first third. Call A (Fig. 45 A–F and 42D). We examined 14 recordings, a total of 31 minutes, with ca. 820 calls from 50 males. Only some of these calls were measured (see Table 2). Call duration varies from 0.691 to 0.835 s. The envelope of the call is variable (Fig. 45A, C, D). In most calls, the limits between the call rise, sustain, and call fall are not clear. Calls usually have a short segment with very low amplitude at the beginning of the call, separated from the rest of the call by an abrupt change in amplitude. The shape of the call rise and fall is usually exponential. The sustain is irregular, usually composed of a shallow or deep valley (i.e., with a concave shape; Fig. 45D). The amplitude peak is often at around the middle or after one third of the call duration. The envelope varies from elliptic (Fig. 45A, D) to triangular (pointed right; Fig. 45C). Due to the concave shape of the sustain, the triangular shape of some calls resembles an arrow. More than 50 % of the call energy is concentrated in 27 % of the call duration around the amplitude peak. There is no PAM in the call. The call has a harmonic series (Fig. 42D). The fundamental frequency is ca. 1020 Hz and the first six harmonics are generally emphasized. The wave periods are regular and harmonics are clear throughout the call. The dominant frequency varies from ca. 950 to 1030 Hz (Fig. 42D). The first harmonic is the dominant (Fig. 42D, 45B, E, F). There is a clear shift in relative energy among bands. Although, usually, there is no shift in the dominant frequency, the higher bands get more energy towards the end of the call (Fig. 42D). Most of the call energy is between 850 and 1150 Hz (one harmonic). The call has a general downward FM (45B, E, F). Additionally, calls have an up-downward FM in the first third of the call duration, leading to arc-shaped bands in this part of the call, and a short upward FM at the end (45B, E, F). The general downward FM and the initial updownward FM result in S-shaped harmonics when considering the entire call. There is no PFM., Published as part of Hepp, Fábio & Pombal, José P., 2020, Review of bioacoustical traits in the genus Physalaemus Fitzinger, 1826 (Anura: Leptodactylidae: Leiuperinae), pp. 1-106 in Zootaxa 4725 (1) on page 76, DOI: 10.11646/zootaxa.4725.1.1, http://zenodo.org/record/3612996

Physalaemus biligonigerus (Cope, 1861) We found a single call type for the species, referred to as call A. The call is composed of a single harmonic note. It has a general downward FM throughout the call but with an up-downward FM segment in the first fifth of the call. Call A (Fig. 46 A–F and 42E). We examined 29 recordings, a total of 65 minutes, with ca. 2140 calls from 105 males. Only some of these calls were measured (see Table 2). Call duration varies from 0.546 to 0.640 s. The envelope of the call is variable (Fig. 46A, C, D). In most calls, the limits between the call rise, sustain, and call fall are not clear. The ratio between call rise and fall duration, and their shapes, are highly variable. Most calls have rise and fall similar in duration, or the former longer than the fall. The shape of the envelopes varies from exponential or linear to logarithmic. The call rise can have two consecutive exponential parts, the first shorter than the second. The sustain is usually irregular, composed of shallow valleys and small peaks (Fig. 46A, C, D). In some calls, the call rise remains with very low amplitude until the limit with the sustain, where the amplitude increases abruptly (Fig. 46D). The amplitude peak is at around the end of the first third or two thirds of the call duration. The envelope varies from rectangular (Fig. 46C) to triangular (pointed left or right; Fig. 46D, A, respectively). Due to the asymmetry of some triangular envelopes, the shape resembles an arrow. More than 50 % of the call energy is concentrated in 30 % of the call duration around the amplitude peak. There is no PAM in the call. The call has a harmonic series (Fig. 42E). The fundamental frequency is ca. 570 Hz and approximately the first eight harmonics are emphasized. The wave periods are regular and harmonics are clear throughout the call. The dominant frequency varies from ca. 600 to 650 Hz (Fig. 46B). The dominant harmonic varies from the first to the sixth (except the second), but it is usually the first (Fig. 42E, 46B, E, F). There is a clear shift in relative energy between the bands; the dominant frequency gets higher toward the end of the call, starting at the first harmonic, moving to the fifth, and ending at the sixth; thenceforth it decreases, ending at the third harmonic (sometimes skipping the fourth harmonic; Fig. 42E, 46B, E, F). Most of the call energy is between 450 and 2950 Hz (four to six harmonics). The call has a general downward FM (Fig. 46B, E, F). Additionally, the calls have an up-downward FM in the first fifth of the call duration, leading to a arc-shaped bands in this part of the call, and a short downward FM at the end (Fig. 46B, E, F). The general downward FM and the initial up-downward FM result in S-shaped harmonics when considering the entire call. There is no PFM., Published as part of Hepp, Fábio & Pombal, José P., 2020, Review of bioacoustical traits in the genus Physalaemus Fitzinger, 1826 (Anura: Leptodactylidae: Leiuperinae), pp. 1-106 in Zootaxa 4725 (1) on page 77, DOI: 10.11646/zootaxa.4725.1.1, http://zenodo.org/record/3612996

Physalaemus marmoratus (Reinhardt & Lütken, 1862) We found a single call type for the species, referred to as call A. The call is composed of a single harmonic note. It has a general downward FM, with an up-downward FM segment in the first third of the call. Call A (Fig. 47 A–J and 42F). We examined 15 recordings, a total of 31 minutes, with ca. 1100 calls from 44 males. Only some of these calls were measured (see Table 2). Call duration varies from 0.614 to 0.938 s. The envelope of the call is highly variable (Fig. 47A, C–F). In most calls, the limits between the call rise, sustain, and call fall are not clear. The ratio between durations of call rise and fall, and their shapes, are highly variable. The sustain is usually irregular, composed of shallow valleys and small peaks (Fig. 47A, C–F). In some calls the rise remains with very low amplitude until the limit with the sustain, where the amplitude increases abruptly (Fig. 47D). In other calls, the call fall has this same pattern, with an abruptly amplitude decrease after the sustain and thenceforth with low and constant amplitude until the end of the call (Fig. 47C). The amplitude peak is usually at around the end of the first third of the call duration. The envelope of the calls varies from rectangular (Fig. 47E, F) to triangular (pointed left or right; Fig. 47D, A, respectively). Due to the asymmetry of some triangular envelopes, the shape resembles an arrow. More than 50 % of the call energy is concentrated in 29 % of the call duration around the amplitude peak. There is no PAM in the call. The call has a harmonic series (Fig. 42F). The fundamental frequency is ca. 510 Hz and the first six harmonics are generally emphasized. The wave periods are regular and harmonics are clear throughout the call. The dominant frequency is ca. 500 Hz (Fig. 47B). The first is dominant harmonic (Fig. 42F). There is a clear shift in relative energy between bands; although there is no shift in dominant frequency, the higher bands get more energy toward the end of the call (Fig. 47 G–J). Most of the call energy is between 400 and 2100 Hz (three to five harmonics). The call has a general downward FM (Fig. 47B, G–J). Additionally, the calls have an up-downward FM in the first third of the call duration, leading to slightly arc-shaped bands in this part of the call, and short downward FM at the end (Fig. 47B, G–J). The general downward FM and the initial up-downward FM result in S-shaped harmonics when considering the entire call. There is no PFM., Published as part of Hepp, Fábio & Pombal, José P., 2020, Review of bioacoustical traits in the genus Physalaemus Fitzinger, 1826 (Anura: Leptodactylidae: Leiuperinae), pp. 1-106 in Zootaxa 4725 (1) on page 78, DOI: 10.11646/zootaxa.4725.1.1, http://zenodo.org/record/3612996

Physalaemus signifer (Girard, 1853) We found two different calls, referred to as call A and B. Calls B were recorded in agonistic contexts (M. Bilate, personal communication). Calls A and B are composed of harmonics and a single note each. Call B tends to be longer and with lower fundamental frequency than Call A. Calls B have strong FM segments and nonlinear regimes, such as deterministic chaos and subharmonics. Call A (Fig. 17 A–L and 13D). We examined 68 recordings, a total of 213 minutes, with ca. 5800 calls from 135 males. Only some of these calls were measured (see Table 2). Call duration varies from 0.453 to 0.579 s. Call rise and fall are short and usually similar to each other in duration. In some calls, the rise is longer than the fall (Fig. 17A, C, D, F). Both can have a linear or exponential shape. There is a long sustain (Fig. 17A, C, D, E, F, G). Usually it has a convex shape it can be quite irregular with periods of concave (amplitude valley) and convex shapes (Fig. 17E, D, respectively). In some calls the rise and the sustain can be fused. The amplitude peak of the call is at around the end of the first four fifths of the call duration (Fig. 17A, C, E, F). The envelope of the call can be classified as elliptic (Fig. 17D), rectangular (Fig. 17F, G), or triangular (pointed left; Fig. 17A, C, E) depending on the shape of the sustain and position of the amplitude peak in the call. More than 50 % of the call energy is concentrated in 34 % of the call duration around the amplitude peak. Some calls have a slight PAM (there is no silence interval between the amplitude peaks; Fig. 17 C, E, G). The rate of the PAM is ca. 50 Hz, forming ca. 22 cycles throughout the call. The cycle rise and fall are similar, with amplitude peak at the middle of the cycle. The call has a harmonic series (Fig. 13D). The fundamental frequency is ca. 280 Hz and this band can be present with low energy or absent in the audiospectrograms. The wave periods are regular and the harmonics are clear throughout the call. Subharmonics are present at the very end of some calls. The dominant frequency varies from ca. 860 to 1550 Hz (Fig. 17B). The dominant harmonic varies from the third to the fifth harmonic, but it is usually the third. There is no clear shift in the relative energy between the bands throughout the call. Most of the call energy is between 700 and 1000 Hz (two harmonics). Usually, the call has a general upward FM with a short downward FM at the end (Fig. 17B, H, I, J, K, L). Some calls have an up-downward FM at the beginning, yielding arc-shaped bands in this part of the call (Fig. 17L). Additionally, there is PFM throughout the call, which is directly proportional to the synchronic PAM (Fig. 17C, E, G, H, J, L). Call B (Fig. 17 M–R and 16B). We examined two recordings, a total of eight minutes, with 64 calls from four males. Only some of these calls were measured (see Table 2). Call duration varies from 0.883 to 1.355 s. Call rise and fall are short and usually similar in duration. In some calls, the rise is longer than the fall. Both can have a linear or exponential shape. There is a long sustain, usually very irregular, with several amplitude peaks (Fig. 17M, O, P). The highest amplitude peak is at around the end of the first nine tenths of the call duration (see below; Fig. 17M, O, P). The envelope of the call can be classified as elliptic, rectangular, or triangular (pointed left; Fig. 17M, O, P) depending on the shape of the sustain and position of the amplitude peak in the call. More than 50 % of the call energy is concentrated in 40 % of the call duration around the amplitude peak. The call has an irregular PAM (there is no silence interval between the amplitude peaks; Fig. 17M). Amplitude peaks are variable in intensity and some of them can show high amplitude. Usually, that is the case of the last peak, where the amplitude peak of the call is. The rate of the PAM is ca. 19 Hz even though highly variable, forming ca. 14 peaks throughout the call. The cycle ride and fall are usually similar, with amplitude peak at the middle of the cycle. The call has a harmonic series (Fig. 16B). The fundamental frequency is ca. 230 Hz and this band can be present with low energy or absent in the audiospectrograms. Usually the wave periods are regular and harmonics are clear throughout the call. However, some parts of the call can have poorly distinguished harmonics, with considerably deterministic chaos due to the high irregularity of the wave periods (Fig. 17Q). Sudden jumps of the fundamental frequency can be present (usually at the end of the call; Fig. 17Q). Moreover, some calls show subharmonics, usually at their ends (Fig. 17N). The dominant frequency varies from ca. 840 to 950 Hz (Fig. 17N). The dominant harmonic varies from the third to the fifth harmonic, but it is usually the fourth. There is no clear shift in the relative energy between the bands throughout the call. Most of the call energy is between 600 and 1200 Hz (two or three harmonics). The call has a general upward FM with a short downward FM at the end (Fig. 17N, Q, R). Additionally, there is a remarkable PFM throughout the call, which is directly proportional to the synchronic PAM where the latter is present (Fig. 17 M–R)., Published as part of Hepp, Fábio & Pombal, José P., 2020, Review of bioacoustical traits in the genus Physalaemus Fitzinger, 1826 (Anura: Leptodactylidae: Leiuperinae), pp. 1-106 in Zootaxa 4725 (1) on page 47, DOI: 10.11646/zootaxa.4725.1.1, http://zenodo.org/record/3612996

Physalaemus obtectus Bokermann, 1966 We found a single call type for the species, referred to as call A. The call is composed of a sequence of pulses. There are clear harmonics, however deterministic-chaos regimes can be present at the beginning of the pulses where jumps of the fundamental frequency are common. Call A (Fig. 10 A–J and 4G). We examined seven recordings, a total of eight minutes, with ca. 280 calls from eight males. Only some of these calls were measured (see Table 2). Call duration varies from 0.415 to 0.553 s. Usually, call rise and fall durations are similar, both resembling logarithmic shape; the amplitude peak is at around (usually just before) the middle of the call duration (Fig. 10A, D, E, F). However, some calls have an exponential or linear rise, followed by a long sustain and an abrupt fall (Fig. 10C). In calls with rise and fall similar in slope and duration, the envelope of the call is symmetric (Fig. 10A, D, E, F), whereas in calls with sustain the envelope is rectangular or triangular (pointed left; Fig. 10C). More than 50 % of the call energy is concentrated in 35 % of the call duration around the amplitude peak. The call has pulse-PAM (with silence intervals present between pulses; Fig. 10 A–J). The rate of the PAM is ca. 9 Hz, forming ca. four pulses throughout the call. The pulses of the first half of the call have rise similar to fall and the amplitude peak is at the middle of the pulse. On the other hand, the pulses of the second half have rises sharper and shorter than falls, which are more gradual, with amplitude peaks at the beginning of the pulses (Fig. 10A, D, E, F). In some calls, the last pulse is clearly shorter than the others (Fig. 10E). There is a long silence interval between the pulses, equivalent to ca. 1.5 times the pulse duration. The call has a harmonic series (Fig. 4G). The fundamental frequency is ca. 380 Hz and this band can be present with low energy or absent in the audiospectrograms. Most of the wave periods are regular and the harmonics are clear throughout the call. However, subharmoncis, deterministic chaos, and jumps of the fundamental frequency are observed at the beginning and end of the pulses (Fig. 10H, I). At the same parts of the pulse, the low fundamental frequency can lead to the wave peaks to be shown as broadband clicks (instantaneously high sound-pressure effect; see Littlejohn 2001) in audiospectrograms at broad filter bandwidths. The dominant frequency varies from ca. 1210 to 1230 Hz (Fig. 10B). The dominant harmonic varies from the third to the seventh, but it is usually the third or fourth. There is no clear shift in the relative energy among the bands throughout the call. Most of the call energy is between 800 and 1600 Hz (often, two or three harmonics). The frequency bands have a general upward FM throughout the call (Fig. 10B, G, I, J). Additionally, there is PFM throughout the call, which is directly proportional to the synchronic pulse-PAM, i.e., each pulse has an up-downward FM (Fig. 10A, B, D–F, H–J)., Published as part of Hepp, Fábio & Pombal, José P., 2020, Review of bioacoustical traits in the genus Physalaemus Fitzinger, 1826 (Anura: Leptodactylidae: Leiuperinae), pp. 1-106 in Zootaxa 4725 (1) on page 38, DOI: 10.11646/zootaxa.4725.1.1, http://zenodo.org/record/3612996, {"references":["Littlejohn, M. J. (2001) Pattern of differentiation in temporal properties of Acoustic signals of anurans. In: Michael, J. R. (Ed.), Anuran Communication. Smithsonian Institution Press, Washington, pp. 102 - 120."]}

Physalaemus spiniger (Miranda-Ribeiro, 1926) We found two different calls, referred to as call A and B. Calls A and B are composed of harmonics and a single note each. Call B has two very different components. The first component is similar to call A, whereas the second one is composed of nonlinear regimes such as deterministic chaos and subharmonics. Call A (Fig. 14 A–L and 13B). We examined 16 recordings, a total of 43 minutes, with ca. 800 calls from 12 males. Only some of these calls were measured (see Table 2). Call duration varies from 0.221 to 0.492 s. The envelope of the call is highly variable (Fig. 14A, C–G). Often, the call rise and fall are similar in duration and vary from linear to exponential shape; there is a long regular sustain or a shallow valley connecting them (Fig. 14A, C). The amplitude peak of the calls is at around the end of the first three fourths of the call duration. The envelope of the call can be elliptic (Fig. 14D, F), rectangular (Fig. 14A, C), or triangular (Fig. 14E, G) depending on the shape of the sustain. There can be a short and deep amplitude valley at the beginning of the call (Fig. 14G). More than 50 % of the call energy is concentrated in 35 % of the call duration around the amplitude peak. The call has no PAM. The call has a harmonic series (Fig. 13B). The fundamental frequency is ca. 380 Hz and this band can be present with low energy or absent in the audiospectrograms. The wave periods are regular and the harmonics are clear throughout the call. The dominant frequency of the call varies from 1120 to 1981 Hz (Fig. 14B). The dominant harmonic varies from the second to the sixth, but it is usually the fourth. There is no clear shift in the relative energy between the bands throughout the call. Most of the call energy is between 1100 and 1700 Hz (ca. two or three harmonics). The call usually has a general upward FM (Fig. 14I, J, K), a few calls have downward FM (Fig. 14B, H, L). Additionally, many calls have a rapid up-downward FM at their outset, forming arc-shaped bands in this part of the call and a short downward FM at the end (Fig. 14B, J, K, L). In calls with short amplitude valleys the bands have down-upward FM, synchronically to the valley (Fig. 14L). Some calls showed a slight PFM (Fig. 14I, J). Call B (Fig. 14 M–P and 6F). We examined two recordings, a total of five minutes, with seven calls from three males. Most of these calls were measured (see Table 2). Call duration varies from 0.376 to 0.512 s. The call rise and fall are short and similar in duration; there is a long irregular sustain (Fig. 14M, O). The amplitude peak is at around the middle of the call duration. Shallow amplitude valleys are present yielding secondary amplitude peaks at the beginning and end of the call. The envelope of the call is somewhat rectangular (Fig. 14M, O). More than 50 % of the call energy is concentrated in 43 % of the call duration around the amplitude peak. The call has no PAM. The call has a harmonic series (Fig. 6F). The call has two different components. The traits of the first component are similar to those of the call A, whereas in the second component, the harmonics can be not well defined due to deterministic chaos (high irregularity of the wave periods; Fig. 14N, P). Subharmonics are common in the second component of the call (Fig. 14N, P). The fundamental frequency is ca. 290 Hz. Although the amplitude often decreases slightly at the transition between components, their limits can be unclear in oscillograms (Fig. 14M). The dominant frequency of the call varies from 1163 to 1723 Hz (Fig. 14N). The dominant harmonic of the second component varies from the third to the 12 th, but it is usually the fourth, fifth, or sixth. There is a slight increase in the relative energy of the higher harmonics throughout the call (Fig. 14N, P). Most of the second component’s energy is between 1200 and 2000 Hz (ca. three harmonics). The general FM is usually downward, with no additional FM at the end (Fig. 14N, P). Some calls have an up-downward FM at the beginning, yielding arc-shaped bands in this part of the call (Fig. 14N). In highly variable calls, and mainly along the second component, there can be several parts with irregular up and downward FM mirroring the AM direction (Fig. 14N, P)., Published as part of Hepp, Fábio & Pombal, José P., 2020, Review of bioacoustical traits in the genus Physalaemus Fitzinger, 1826 (Anura: Leptodactylidae: Leiuperinae), pp. 1-106 in Zootaxa 4725 (1) on page 43, DOI: 10.11646/zootaxa.4725.1.1, http://zenodo.org/record/3612996

Physalaemus gracilis (Boulenger, 1883) We found a single call type for the species, referred to as call A. The call is composed of a single harmonic note, with general downward FM, but with an up-downward FM segment in the first sixth of the call. Call A (Fig. 50 A–T and 52A). We examined 40 recordings, a total of 158 minutes, with ca. 2480 calls from 106 males. Only some of these calls were measured (see Table 2). Call duration varies from 0.451 to 0.565 s. The envelope of the call is variable (Fig. 50A, C–G, M–P). In most calls, rise and fall are similar in duration and shape (exponential) and the sustain is long. Some sustains are regular (Fig. 50A, G, M, N, O) and others are irregular, with short and shallow valleys, mainly at the beginning of the segment (Fig. 50D, F, P). In several calls, the sustain has a convex shape and the limits between the call rise, sustain, and call fall are not clear. Usually, the envelope is divided into two parts with different amplitude levels (Fig. 50C). The amplitude peak is usually at the end of the first seven tenths of the call duration. The envelope varies from elliptic or rectangular (Fig. 50A, D, F, G, M, N, O) to triangular (pointed left; Fig. 50C, E, P). Due to the asymmetry of some triangular envelopes, the shape resembles an arrow. More than 50 % of the call energy is concentrated in 32 % of the call duration around the amplitude peak. There is no PAM in the call. The call has a harmonic series (Fig. 52A). The fundamental frequency is ca. 510 Hz and approximately the first seven harmonics are emphasized. The wave periods are regular and harmonics are clear throughout the call. Subharmonics (f 0 1/2 and f 0 1/3) are common at the beginning and middle of the call (Fig. 50I, L, Q, R). The dominant frequency varies from ca. 2110 to 2760 Hz (Fig. 50B). The dominant harmonic varies from the first to the seventh (except the second), but it is usually the fourth, fifth, or sixth (Fig. 52A). There is a clear shift in relative energy between the bands; the dominant frequency gets higher toward the end of the call, starting at the first harmonic, moving to the third, fourth, fifth and sixth, and ending at the sixth or seventh; thenceforth, the dominant frequency decreases in some calls, moving to the fourth harmonic (Fig. 52A, 50B). Most of the call energy is between 950 and 3050 Hz (four to five harmonics). The call has a general downward FM (Fig. 50B, H–L, Q–T). Additionally, calls have an up-downward FM in the first sixth of the call duration, leading to slightly arc-shaped bands in this part of the call, and a short downward FM at the end (Fig. 50B, H–L, Q–T). The general downward FM and the initial up-downward FM result in S-shaped harmonics when considering the entire call. Some calls have a slight PFM (Fig. 50H, I, L)., Published as part of Hepp, Fábio & Pombal, José P., 2020, Review of bioacoustical traits in the genus Physalaemus Fitzinger, 1826 (Anura: Leptodactylidae: Leiuperinae), pp. 1-106 in Zootaxa 4725 (1) on page 81, DOI: 10.11646/zootaxa.4725.1.1, http://zenodo.org/record/3612996

Physalaemus irroratus Cruz, Nascimento & Feio, 2007 We found a single call type for the species, referred to as call A. The call is composed of a sequence of pulses. There are clear harmonics, and some pulses have PFM, which is synchronic and directly proportional to the slight PAM. Call A (Fig. 11 A–F and 4H). We examined two recordings, a total of one minute, with 29 calls from two males. Most of these calls were measured (see Table 2). Call duration varies from 0.489 to 0.954 s. The call rise and fall durations are similar; both are usually linear-shaped. In some calls, the rise and/or fall can be more abrupt and have a logarithmic shape. The amplitude peak is at around the middle of the call duration, except in calls with very few pulses (e.g., two pulses). Since both rise and fall are similar in slope and duration, the envelope of the call is fairly elliptic (Fig. 11A, C). More than 50 % of the call energy is concentrated in 38 % of the call duration around the amplitude peak. The call has a strong PAM (with silence intervals present between pulses; Fig. 11 A–F). The rate of the PAM is ca. 6 Hz, forming ca. four pulses throughout the call. Usually, pulses have an abrupt rise, shorter than fall, which is more gradual, with amplitude peak at the beginning of the pulse (Fig. 11A, C, D). However, the first pulses of the longer calls have rises similar to falls and the amplitude peak is at the middle of the pulse. In some calls, the last pulse is clearly shorter than the others. There is a long silence interval between pulses, ca. 5.5 times the pulse duration (Fig. 11A, C, D). Some pulses have an internal slight PAM. The call has a harmonic series (Fig. 4H). The fundamental frequency is ca. 400 Hz and this band can be present with low energy or absent in the audiospectrograms. The wave periods are regular and then the harmonics are clear throughout the call. However, subharmonics, deterministic chaos, and jumps of the fundamental frequency are observed at the beginning and at the end of the pulses, or even in the entire pulse (usually the last one). Short pulses can be shown as broadband clicks (instantaneously high sound-pressure effect; see Littlejohn 2001) in audiospectrograms at broad filter bandwidth (first pulse in Fig. 11A). The dominant frequency varies from ca. 1250 to 1720 Hz (Fig. 11B). The dominant harmonic varies from the second to the ninth, but it is usually the third or fourth. There is no clear shift in the relative energy among the bands throughout the call. Most of the call energy is between 1300 and 2000 Hz (often, three harmonics). The frequency bands have a general upward FM throughout the call (Fig. 11B). There is PFM throughout the call, which is directly proportional to the synchronic pulse-PAM, i.e., each pulse has an up-downward FM (Fig. 11F). Additionally, another PFM is present within some pulses and it is directly proportional to the synchronic slight PAM within the pulses., Published as part of Hepp, Fábio & Pombal, José P., 2020, Review of bioacoustical traits in the genus Physalaemus Fitzinger, 1826 (Anura: Leptodactylidae: Leiuperinae), pp. 1-106 in Zootaxa 4725 (1) on page 39, DOI: 10.11646/zootaxa.4725.1.1, http://zenodo.org/record/3612996, {"references":["Cruz, C. A. G., Nascimento, L. B. & Feio, R. N. (2007) A new species of the genus Physalaemus Fitzinger, 1826 (Anura, Leiuperidae) from Southeastern Brazil. Amphibia-Reptilia, 28, 457 - 465. https: // doi. org / 10.1163 / 156853807782152444","Littlejohn, M. J. (2001) Pattern of differentiation in temporal properties of Acoustic signals of anurans. In: Michael, J. R. (Ed.), Anuran Communication. Smithsonian Institution Press, Washington, pp. 102 - 120."]}

Physalaemus nanus (Boulenger, 1888) We found three different calls, referred to as call A, B, and C (Fig. 12). Calls B and C were common in recordings in which several males were active and calling close to each other. Calls A and B are composed of harmonics and a single note each. Call B is shorter than Call A with a lower fundamental frequency, irregular FM segments, and absence of pulse-PAM. Call C is composed of two notes, the first and the second notes are similar to those of calls A and B, respectively. Call A (Fig. 12 E–H and 13A). We examined 20 recordings, a total of 77 minutes, with ca. 3500 calls from 33 males. Only some of these calls were measured (see Table 2). Call duration varies from 0.178 to 0.218 s. The call envelope is variable; however, calls often have rise, a regular sustain (or shallow valley), and falls sections. Call rise and fall are usually gradual and linear but they can have different durations, being long or abrupt. The amplitude peak of the calls measured here is at around the end of the first fourth of the call duration (Fig. 12A, C, D, E). The envelope of the call can be elliptic (Fig. 12A, D), rectangular (Fig. 12E), or triangular (Fig. 12C). More than 50 % of the energy is concentrated in 42 % of the call duration around the amplitude peak. This call has a strong PAM (with silence intervals present between pulses; Fig. 12 A–H). The rate of the PAM is ca. 28 Hz, forming ca. five pulses throughout the call. The envelope of the pulses is also highly variable; however, the middle pulses tend to have amplitude peak at the middle of the pulse with similar rise and fall. Often, the first pulse has very little amplitude and the last pulse is the longest one (Fig. 12C, D, F, G). Silence intervals are present between pulses, slightly shorter than pulse duration (Fig. 12 A–H). Some pulses have a down-upward AM at the middle of their durations, yielding two amplitude peaks per pulse. The call has a harmonic series (Fig. 13A). The fundamental frequency varies from 620 to 1100 Hz and the band can be present with low energy or absent in the audiospectrograms. The wave periods are regular and then the harmonics are clear throughout the call. Subharmonics can be present at the beginning and end of the pulses and jumps of the fundamental frequency can happen at the end of the call (fourth pulse in Fig. 12B). The dominant frequency varies from ca. 2240 to 2540 Hz (Fig. 12B). The dominant harmonic varies from the second to the fourth one, but it is usually the second. There is no clear shift in the relative energy among the bands throughout the call. Most of the energy is concentrated between 1800 and 2800 Hz (ca. two harmonics). The frequency bands have a general and slight downward FM throughout the call (Fig. 12B). Additionally, there is PFM throughout the call, which is directly proportional to the synchronic pulse-PAM (Fig. 12 A–H). Call B (Fig. 12 I–N and 6D). We examined five recordings, a total of 27 minutes, with ca. 40 calls from 13 males. Only some of these calls were measured (see Table 2). Call duration varies from 0.027 to 0.090 s. Often, the call rise is longer than the fall, both exponential; there is a long regular sustain (or shallow valley) between them. The amplitude peak is at around the end of the first three fourths of the call duration (Fig. 12I, K, L). The envelope of the call varied from rectangular (Fig. 12K) to triangular (pointed left; Fig. 12I, L). More than 50 % of the energy is concentrated in 31 % of the component duration around the amplitude peak. This call has no PAM. The call has a harmonic series (Fig. 6D). The fundamental frequency is ca. 300 Hz and this band can be present with low energy or absent in the audiospectrograms. Usually, the wave periods are regular and then the harmonics are clear throughout the call. However, harmonics are not very clear with considerably deterministic chaos in some parts of the call (Fig. 12N). Sudden jumps of the fundamental frequency can be present (usually at the end of the call). Moreover, some calls show subharmonics (Fig. 12N). The dominant frequency varies from ca. 1680 to 1850 Hz (Fig. 12J). The dominant harmonic varies from the seventh to the 41 st, but it is usually the ninth or tenth. There is no clear shift in the relative energy among the bands throughout the call. Most of the energy is concentrated between 1300 and 2200 Hz (three or four harmonics). The frequency bands can have a general down or upward FM throughout the call with either short down or upward FM at the end (Fig. 12J, M, N). Some calls have no clear general FM. Additionally, some calls have a subtle PFM throughout the call (Fig. 12J, M, N). Call C (Fig. 12 O–T and 6E). We examined 16 recordings, a total of 60 minutes, with ca. 200 calls from 28 males. Only some of these calls were measured (see Table 2). Calls are composed of two notes, the first and the second are similar to those of calls A and B, respectively. Call duration varies from 0.188 to 0.311 s. The amplitude, temporal, and spectral traits of the components are similar to those described above. However, the first note can have more pulses and the envelope of the second note has steeper rise and fall in call C (Fig. 12O, Q, R). Although the amplitude decreases at the transition between notes, their limits are not clear (Fig. 12Q, R). At this transition, there is a decrease in the fundamental frequency and wave peaks emitted at low repetition rates (e.g., 90 Hz) are shown as clicks (instantaneously high sound-pressure effect; Fig. 12P) in audiospectrograms at broad filter bandwidths (e.g., above 100 Hz). The bands are observed in audiospectrograms at narrow filter bandwidth (e.g., below 90 Hz). This rate gets faster until the beginning of the center of the second note. In C calls, the harmonics of the second note usually have a general upward FM with a short downward FM at the end of the call (Fig. 12P but see Fig. 12S, T)., Published as part of Hepp, Fábio & Pombal, José P., 2020, Review of bioacoustical traits in the genus Physalaemus Fitzinger, 1826 (Anura: Leptodactylidae: Leiuperinae), pp. 1-106 in Zootaxa 4725 (1) on pages 40-43, DOI: 10.11646/zootaxa.4725.1.1, http://zenodo.org/record/3612996

Physalaemus santafecinus Barrio, 1965 We found a single call type for the species, referred to as call A. The call is composed of a single harmonic note. It has a general downward FM, with an up-downward FM segment in the first third of the call. Call A (Fig. 48 A–D and 42G). We examined two recordings, a total of three minutes, with 61 calls from two males. Only some of these calls were measured (see Table 2). Call duration varies from 0.330 to 0.375 s. The envelope of the call is variable. In most calls, the rise and fall are similar in duration and shape (exponential). In some calls, the limits between call rise, sustain, and call fall are not clear, with linear or logarithmic-shaped rise and fall and the sustain with a convex shape (calls with elliptic envelope; Fig. 48C). The amplitude peak is usually at around the end of the first seven tenths of the call duration or at the middle of the call. The envelope varies from elliptic (Fig. 48C) to triangular (pointed left; Fig. 48A). Due to the concave shape of the sustain, the triangular envelope of some calls resembles an arrow (Fig. 48A). More than 50 % of the call energy is concentrated in 36 % of the call duration around the amplitude peak. There is no PAM in the call. The call has a harmonic series (Fig. 42G). The fundamental frequency is ca. 490 Hz and approximately the first seven harmonics are emphasized. The wave periods are regular and harmonics are clear throughout the call. The dominant frequency varies from ca. 474 to 2627 Hz (fig. 48B). The dominant harmonic varies from the first to sixth (rarely the second harmonic), but it is usually the first (Fig. 42G, 48B, D). There is a clear shift in relative energy between the bands; the dominant frequency gets higher toward the end of the call, starting at the first harmonic, moving to the fifth and ending at the sixth; thenceforth it decreases, usually skipping the fifth and ending at the third or second harmonic (Fig. 42G, 48B, D). Most of the call energy is between 450 and 2650 Hz (four to six harmonics). The call has a general downward FM (Fig. 48B, D). Additionally, calls have an up-downward FM in the first third of the call duration, leading to slightly arc-shaped bands in this part of the call (Fig. 48B), and a short downward FM at the end (Fig. 48B, D). The general downward FM and the initial up-downward FM result in S-shaped harmonics when considering the entire call. There is no PFM., Published as part of Hepp, Fábio & Pombal, José P., 2020, Review of bioacoustical traits in the genus Physalaemus Fitzinger, 1826 (Anura: Leptodactylidae: Leiuperinae), pp. 1-106 in Zootaxa 4725 (1) on page 79, DOI: 10.11646/zootaxa.4725.1.1, http://zenodo.org/record/3612996, {"references":["Barrio, A. (1965) El genero Physalaemus (Anura, Leptodactylidae) en la Argentina. Physis, 25 (70), 421 - 448."]}

Physalaemus orophilus Cassini, Cruz & Caramaschi, 2010 We found a single call type for the species, referred to as call A. The call has long duration and is composed of a single harmonic note with a sequence of pulses with interpulse silence intervals. The bands have a general upward FM but with a subtle downward FM at the end, yielding slightly arc-shaped bands in the audiospectrogram of some calls when considering the entire call duration. Call A (Fig. 41 A–H and 42A). We examined 10 recordings, a total of 26 minutes, with ca. 270 calls from 16 males. Only some of these calls were measured (see Table 2). Call duration varies from 3.724 to 5.432 s. Call rise and fall are very short and similar to each other in duration. There is a long sustain. This segment is usually regular and almost flat (Fig. 41A, D), but convex in some calls (Fig. 41E). The amplitude peak of these calls is at the end of the first two thirds of the call duration. The envelope of the call varies from elliptic (Fig. 41E) to rectangular (Fig. 41A, D). More than 50 % of the call energy is concentrated in 43 % of the call duration around the amplitude peak. The call has a strong PAM (silence intervals are present between peaks; Fig. 41 A–H). The rate of the PAM is ca. 11 Hz, forming ca. 57 pulses throughout the call. The pulse rise is longer than fall, with amplitude peak of the pulse at ca. two thirds of the pulse duration. The amplitude peak of the last pulse is at the beginning or middle of the pulse (Fig. 41C). Duration of silence intervals is similar to pulse duration. The last pulse is usually the longest (ca. 1.5 times longer than the other pulses). In some calls, the last pulse is the shortest. The call has a harmonic series (Fig. 42A). The fundamental frequency is ca. 290 Hz. The first five harmonics are usually absent in audiospectrograms (Fig. 41B). There are ca. four adjacent emphasized harmonics. The wave periods are regular and harmonics are clear throughout the call. The dominant frequency varies from ca. 2630 to 2910 Hz (Fig. 41B). The dominant harmonic varies from the seven to the 29 th, but it is usually the ninth (Fig. 42A). There is no shift in the relative energy between the bands throughout the call. Most of the call energy is between 2500 and 3350 Hz (three harmonics; Fig. 41F). The call has a slight general upward FM and a short downward FM at the end, leading to arc-shaped bands in audiospectrograms when considering the entire call (Fig. 41B, G, H). Additionally, there can be a slight PFM throughout the call, which is directly proportional to the synchronic pulse-PAM, i.e. up-downward FM in each pulse., Published as part of Hepp, Fábio & Pombal, José P., 2020, Review of bioacoustical traits in the genus Physalaemus Fitzinger, 1826 (Anura: Leptodactylidae: Leiuperinae), pp. 1-106 in Zootaxa 4725 (1) on page 72, DOI: 10.11646/zootaxa.4725.1.1, http://zenodo.org/record/3612996, {"references":["Cassini, C. S., Cruz, C. A. G. & Caramaschi, U. (2010) Taxonomic review of Physalaemus olfersii (Lichtenstein & Martens, 1856) with revalidation of Physalaemus lateristriga (Steindachner, 1864) and description of two new related species (Anura: Leiuperidae). Zootaxa, 2491 (1), 1 - 33. https: // doi. org / 10.11646 / zootaxa. 2491.1.1"]}

Physalaemus barrioi Bokermann, 1967 We found a single call type for the species, referred to as call A. The call has a single harmonic note with a long duration and general downward FM, with an up-downward FM segment in the first seventh of the call. Call A (Fig. 54 A–L and 52D). We examined eight recordings, a total of 13 minutes, with ca. 70 calls from eight males. Only some of these calls were measured (see Table 2). Call duration varies from 1.323 to 2.038 s. Call rise duration is short and similar to call fall duration; the call rise and fall shapes vary from logarithmic to almost linear or exponential. The sustain is irregular, generally flat (Fig. 54A, E, F) or ascending (Fig. C, D, G). In this latter case, the amplitude gets higher towards the end of the call. There is usually a long shallow valley at the beginning or at the middle of the sustain (Fig. 54A, C, D, E, G). The amplitude peak is usually at the end of the first three fifths of the call duration. The envelope varies from elliptic (Fig. 54F), rectangular (Fig. 54A, E, G) to triangular (usually pointed left; Fig. C, D). More than 50 % of the call energy is concentrated in 39 % of the call duration around the amplitude peak. There is no PAM in the call. The call has a harmonic series (Fig. 52D). The fundamental frequency is ca. 460 Hz and the first seven harmonics are generally emphasized. The wave periods are regular and harmonics are clear throughout the call. The dominant frequency varies from ca. 470 to 2580 Hz. The dominant harmonic is the first or the sixth, but usually the first (Fig. 52D, 54B, H–L). There is a clear shift in relative energy between the bands; the dominant frequency increases towards the end of the call, starting at the first harmonic and moving to the sixth at the very end of the call (Fig. 52D, 54B, H–L). Most of the call energy is between 450 and 2700 Hz (four to six harmonics). The call has a general downward FM. Additionally, the calls have an up-downward FM at the first seventh of call duration, leading to slightly arc-shaped bands in this part of the call, and a short downward FM at the end (Fig. 54B, H–L). The general downward FM and the initial up-downward FM result in S-shaped harmonics when considering the entire call. There is no PFM., Published as part of Hepp, Fábio & Pombal, José P., 2020, Review of bioacoustical traits in the genus Physalaemus Fitzinger, 1826 (Anura: Leptodactylidae: Leiuperinae), pp. 1-106 in Zootaxa 4725 (1) on page 85, DOI: 10.11646/zootaxa.4725.1.1, http://zenodo.org/record/3612996, {"references":["Bokermann, W. C. A. (1967) Tres novas especies de Physalaemus do sudeste brasileiro (Amphibia, Leptodactylidae). Revista Brasileira de Biologia, 27 (2), 135 - 143."]}

Physalaemus albonotatus (Steindachner, 1864) We found a single call type for the species, referred to as call A. The call has a single harmonic note with a slight PAM without silence intervals. It has a gradual downward FM throughout the call. Call A (Fig. 25 A–H and 24B). We examined 14 recordings, a total of 24 minutes, with ca. 330 calls from 26 males. Only some of these calls were measured (see Table 2). Call duration varies from 1.333 to 1.429 s. In most calls the limits between the call rise, sustain, and call fall are not clear (for example, see elliptic envelope in Fig 25A). In calls where they are perceptible, the call rise and fall can be similar in duration with variable shape (linear, exponential, or logarithmic) or call fall is shorter than the rise. In some calls, there is a long regular sustain (Fig. 25E). The amplitude peak of the calls measured is at around the end of the first three fourths of the call duration. The envelope varies from elliptic (Fig. 25A, C, D) to rectangular (Fig. 25E). More than 50 % of the call energy is concentrated in 42 % of the call duration around the amplitude peak. The call has a slight PAM (there is no silence interval between amplitude peaks; Fig. 25A, C, D, E). The rate of the PAM is ca. 25 Hz, forming ca. 35 cycles throughout the call. The cycle rise and fall are similar, with amplitude peak at the middle of the cycle. The call has a harmonic series (Fig. 24B). The fundamental frequency is ca. 530 Hz and approximately the first six harmonics are emphasized. The wave periods are regular and harmonics are clear throughout the call. The dominant frequency varies from ca. 1590 to 2440 Hz (Fig. 25B). The dominant harmonic varies from the first to the sixth, but it is usually the fifth. There is a clear shift in relative energy among the bands; the dominant frequency gets higher toward the end of the call, starting at first harmonic and ending in the fifth or sixth one (Fig. 25B, F, G, H). Most of the call energy is between 450 and 2950 Hz (five to six harmonics). The call has a general downward FM. Additionally the calls have a subtle up-downward FM at the beginning, yielding arc-shaped bands in this part of the call (Fig. 25F, G, H), and a short downward FM at the end (Fig. 25B, F, G, H). The general downward FM and the initial up-downward FM result in S-shaped harmonics when considering the entire call. The call also has a PFM, which is inversely proportional and synchronic to the PAM (Fig. 25A, B, C, E, F, H). In a few calls, the rate of the PAM is very low and so is the number of cycles (Fig. 25D, G). In those calls, the PFM is equally slow and weak (Fig. 25D, G)., Published as part of Hepp, Fábio & Pombal, José P., 2020, Review of bioacoustical traits in the genus Physalaemus Fitzinger, 1826 (Anura: Leptodactylidae: Leiuperinae), pp. 1-106 in Zootaxa 4725 (1) on page 56, DOI: 10.11646/zootaxa.4725.1.1, http://zenodo.org/record/3612996

Physalaemus bokermanni Cardoso & Haddad, 1985 We found two different calls, referred to as call A and B. Calls B were common in recordings in which several males were active and calling at the same night. Calls A and B are composed of harmonics and pulses (i.e., pulse-PAM). Call B has two notes while Call A has only one. The first note of call B is similar to that of call A. The second note of call B is much longer than that of call A and has an envelope with a long and gradual rise. Call A (Fig. 18 A–D and 13E). We examined eight recordings, a total of 12 minutes, with ca. 650 calls from nine males. Only some of these calls were measured (see Table 2). Call duration varies from 0.177 to 0.197 s. The call rise is linear or logarithmic-shaped and longer than the fall, which is usually abrupt and logarithmic-shaped; the amplitude peak is at around the end of the first three fourths of the call duration (Fig. 18A). The envelope of the call is elliptic or triangular (pointed left; Fig. 18A, C). More than 50 % of the energy is concentrated in 41 % of the call duration around the amplitude peak. This call has a strong PAM (with silence intervals present between pulses; Fig. 18 A–D). The rate of the PAM is ca. 35 Hz, forming ca. six pulses throughout the call. The pulse rise is abrupt and much shorter than the fall; the amplitude peak is at the beginning of the pulse (Fig. 18C). The first one or two pulses have much lower amplitude than the others (Fig. 18A). Often, the second and the last pulses are the longest (Fig. 18A, B). Silence intervals are present between pulses, ca. fivefold longer than pulse duration. The first interval is usually much shorter than the others (Fig. 18 A–D). The call has a harmonic series (Fig. 13E). The fundamental frequency is ca. 780 Hz and this band can be present with low energy or absent in the audiospectrograms. The short duration of the pulses makes the bands broad with narrow intervals (Fig. 18B, D). Some pulses can have not very clear harmonics, with considerably deterministic chaos due to the irregularity of the wave periods (Fig. 18B, D). There are jumps of the fundamental frequency between pulses in some calls. The dominant frequency varies from ca. 2840 to 3660 Hz (Fig. 18B). The dominant harmonic varies from the second to the 10 th, but it is usually the fourth. There is no clear shift in the relative energy among the bands throughout the call. Most of the energy is concentrated between 2600 and 3900 Hz (ca. three harmonics). Usually, there is no clear general FM throughout the call, however, in some calls the first two pulses have their energy concentrated in lower frequency bands, making the general FM of the call upward (Fig. 18B). Call B (Fig. 18 E–H and 16C). We examined three recordings, a total of six minutes, with 35 calls from three males. Only some of these calls were measured (see Table 2). Call duration varies from 0.947 to 1.868 s and the call has two different notes. Duration of the second note is ca. 1.0 s. The amplitude, temporal and spectral traits of the first note resemble those of call A, although in call B the first note often has more abrupt rise and fall (Fig. 18E). Usually, there is a silence interval between the notes (Fig. 18E, F). However, in some calls, this interval is perceptible only as a decrease in amplitude. The rise of the second note is logarithmic-shaped and shorter than fall, which is gradual, almost linear; the amplitude peak of the note is at the end of the first tenth of the note duration (Fig. 18E). Due to the very short rise and the long and gradual fall, the envelope of the component B is triangular (pointed right; Fig. 18E). More than 50 % of the energy of the compound call is concentrated in ca. 34 % of the duration around the amplitude peak. Both notes have a strong PAM (there are silence intervals present between pulses; Fig. 18 E–H). The rate of the PAM is similar to that of the call A, ca. 30 Hz, yielding 28 pulses throughout the call. The rate of the PAM is less regular in the second note than in the first one. The pulse rise is abrupt and much shorter than the fall; the amplitude peak is at the beginning of the pulse (Fig. 18G). Some pulses can be twofold longer than the others. At the beginning of the second note the ratio between the silence interpulse-interval and pulse duration is similar to that in component A. The interval becomes longer (pulse duration remains the same) towards the end of the call (i.e., pulse-PAM rate decreases), mainly after the first fourth of the second-note duration (Fig. 18E, G). Spectral traits of the second note are similar to those of call A (Fig. 18F, H; see some quantitative differences in Table 2). There is no general FM in component B (Fig. 18F, H)., Published as part of Hepp, Fábio & Pombal, José P., 2020, Review of bioacoustical traits in the genus Physalaemus Fitzinger, 1826 (Anura: Leptodactylidae: Leiuperinae), pp. 1-106 in Zootaxa 4725 (1) on pages 49-50, DOI: 10.11646/zootaxa.4725.1.1, http://zenodo.org/record/3612996, {"references":["Cardoso, A. J. & Haddad, C. F. (1985) Nova especie de Physalaemus do grupo signiferus (Amphibia, Anura, Leptodactylidae). Revista Brasileira de Biologia, 45, 33 - 37."]}

Physalaemus henselii (Peters, 1872) We found a single call type for the species, referred to as call A. The call is composed of a sequence of pulses (i.e., pulse-PAM). The call is spectrally polymorphic; some calls have pulses with sidebands. Call A (Fig. 23 A–F and 24A). We examined two recordings, a total of four minutes, with ca. 200 calls from six males. Only some of these calls were measured (see Table 2). Call duration varies from 0.289 to 0.493 s. The call rise is usually very abrupt and shorter than the call fall (Fig. 23C), which has an exponential shape. The call rises are longer and more similar to falls. The sustain varies from flat to very steep in shape. The envelope of the call is elliptic, rectangular (Fig. 23C), or triangular (pointed left; Fig. 23A). More than 50 % of the energy is concentrated in 53 % of the call duration around the amplitude peak. The call has a strong PAM (with silence intervals present between pulses; Fig. 23 A–F). The rate of the PAM is ca. 54 Hz, forming ca. 20 pulses throughout the call. The envelope of the pulses is variable; however, the pulse rise is usually shorter than the fall, with amplitude peak at the beginning of the pulse. Silence intervals are present between pulses, with durations slightly longer than pulse duration except between the first pulses, where the intervals are very short or even absent (pulses are juxtaposed; Fig. 23A, C, D). The call has a harmonic series (Fig. 24A). The fundamental frequency is at ca. 1900 Hz and this band is also the dominant frequency (see below). The wave periods are regular and harmonics are clear throughout the call. The call shows an additional frequency series with bands separated by ca. 250 Hz series produced by a PAM present within pulses (Fig. 23E, F). This series is very variable (30 to 550 Hz) and it is not multiple of the harmonic series. Both seem to be independent of each other. Therefore, we called the 250-Hz bands as sidebands. The short duration of the pulses makes the bands broad with narrow intervals. In parts where two pulses are juxtaposed, or at least very close to each other, the wave periods are less regular, the harmonics can be less clear with deterministic chaos (Fig. 23E). The dominant frequency varies from ca. 1690 to 2160 Hz (Fig. 23B). The first harmonic is the dominant. There is no clear shift in the relative energy among the bands throughout the call. Most of the energy is concentrated between 650 and 2600 Hz (one harmonic). Most of the call energy is between 1400 and 2400 Hz. There is usually neither a clear general FM nor other shorter FM segment in the call. Some calls, mainly those with juxtaposed pulses, show a slight PFM following the PAM (see beginning of the call in Fig. 23E)., Published as part of Hepp, Fábio & Pombal, José P., 2020, Review of bioacoustical traits in the genus Physalaemus Fitzinger, 1826 (Anura: Leptodactylidae: Leiuperinae), pp. 1-106 in Zootaxa 4725 (1) on page 55, DOI: 10.11646/zootaxa.4725.1.1, http://zenodo.org/record/3612996

Physalaemus feioi Cassini, Cruz & Caramaschi, 2010 We found a single call type for the species, referred to as call A. The call has long duration and is composed of a single harmonic note with a sequence of pulses with interpulse silence intervals. The bands have a general upward FM and a downward FM at the end, yielding a slight arc shape in the audiospectrogram when considering the entire call. Call A (Fig. 40 A–H and 33H). We examined seven recordings, a total of eight minutes, with 52 calls from nine males. Only some of these calls were measured (see Table 2). Call duration varies from 3.854 to 4.920 s. Call rise and fall are very short and similar to each other in duration. There is a long sustain. This segment is usually regular and almost flat but some calls have sustains with a convex shape (Fig. 40A, D). The amplitude peak is often at the end of the first seven tenths of the call duration. The envelope of the call is rectangular (Fig. 40A, D). More than 50 % of the call energy is concentrated in 45 % of the call duration around the amplitude peak. The call has a strong PAM (there are silence intervals between pulses; Fig. 40A, D). The rate of the PAM is ca. 15 Hz, forming ca. 55 pulses throughout the call. The pulse rise is longer than fall, with amplitude peak of the pulse at ca. two thirds of the pulse duration (Fig. 40C). The amplitude peak of the last pulse is at the beginning or middle of the pulse. Interval durations are similar to pulse duration (Fig. 40C). The last pulse is usually the longest (ca. 1.5 times longer than the other pulses; Fig. 40E). In some calls, the last pulse is the shortest (Fig. 40A). The call has a harmonic series (Fig. 33H). The fundamental frequency is ca. 330 Hz. The first five harmonics are usually absent in the audiospectrogram. There are ca. four adjacent emphasized harmonics. The wave periods are regular and harmonics are clear throughout the call. The dominant frequency varies from ca. 2340 to 2470 Hz. The dominant harmonic varies from the sixth to the 15 th, but it is usually the seventh (Fig. 33H). There is no clear shift in the relative energy between the bands throughout the call. Most of the call energy is between 2100 and 2950 Hz (three harmonics). The call has a general upward FM and a short downward FM at the end, yielding a slight arc shape in the audiospectrogram when considering the entire call (Fig. 40B, G). There is a slight PFM throughout the call, which is directly proportional to the synchronic pulse-PAM, i.e. up-downward FM in each pulse (Fig. 40A, B, C, F). Additionally, there is another PFM, which is perceptible within the pulses (Fig. 40H)., Published as part of Hepp, Fábio & Pombal, José P., 2020, Review of bioacoustical traits in the genus Physalaemus Fitzinger, 1826 (Anura: Leptodactylidae: Leiuperinae), pp. 1-106 in Zootaxa 4725 (1) on page 72, DOI: 10.11646/zootaxa.4725.1.1, http://zenodo.org/record/3612996, {"references":["Cassini, C. S., Cruz, C. A. G. & Caramaschi, U. (2010) Taxonomic review of Physalaemus olfersii (Lichtenstein & Martens, 1856) with revalidation of Physalaemus lateristriga (Steindachner, 1864) and description of two new related species (Anura: Leiuperidae). Zootaxa, 2491 (1), 1 - 33. https: // doi. org / 10.11646 / zootaxa. 2491.1.1"]}

Physalaemus evangelistai Bokermann, 1967 We found a single call type for the species, referred to as call A. The call is composed of a single harmonic note, with a general downward FM and an up-downward FM segment in the first sixth of the call duration. Calls usually have intermediate PAM (with no silence intervals between peaks) and PFM throughout their duration. Call A (Fig. 53 A–F and 52C). We examined eight recordings, a total of 27 minutes, with ca. 340 calls from 20 males. Only some of these calls were measured (see Table 2). Call duration varies from 0.976 to 1.358 s. Call rise duration is very short and similar to call fall duration; the call rise and fall shapes vary from logarithmic to almost linear or exponential. The sustain is flat or gradually ascending (Fig. 53D, C). There is a long shallow valley at the beginning of the sustain (Fig. A, C, D). The amplitude peak is usually at the end of the first seven tenths of the call duration. The envelope varies from rectangular (Fig. 53A, D) to triangular (pointed left; Fig. 53C). More than 50 % of the call energy is concentrated in 36 % of the call duration around the amplitude peak. Some calls show an intermediate PAM only in the final two fourths of the call duration. This PAM yields emphasized cycles (with no silence intervals between peaks; Fig. 53A, C). The rate of the PAM is ca. 9 Hz, forming ca. 10 cycles throughout the call. The cycle rise and fall are similar, with amplitude peak at the middle of the cycle. The call has a harmonic series (Fig. 52C). The fundamental frequency at ca. 540 Hz and approximately the first six harmonics are emphasized. The wave periods are regular and harmonics are clear throughout the call. The dominant frequency varies from ca. 670 to 2910 Hz (Fig. 53B). The dominant harmonic varies from the first to sixth (except the second), but it is usually the first or fourth (Fig. 52C). There is a clear shift in relative energy between the bands; the dominant frequency increases towards the end of the call, starting at the first harmonic, moving to the fourth and fifth, and ending at the sixth; thenceforth, it decreases, ending at the third (Fig. 52C, 53E, F). Most of the call energy is between 550 and 3100 Hz (three to four harmonics). The call has a general downward FM, with a short up-downward FM at the first sixth of the call duration, leading to slightly arc-shaped bands in this part of the call, and a short downward FM at the end (Fig. 53B, E, F). The general downward FM and the initial up-downward FM result in S-shaped harmonics when considering the entire call. Some calls have a PFM during the entire call independent of the PAM. Other calls have PFM inversely proportional and synchronic to the PAM., Published as part of Hepp, Fábio & Pombal, José P., 2020, Review of bioacoustical traits in the genus Physalaemus Fitzinger, 1826 (Anura: Leptodactylidae: Leiuperinae), pp. 1-106 in Zootaxa 4725 (1) on pages 83-84, DOI: 10.11646/zootaxa.4725.1.1, http://zenodo.org/record/3612996, {"references":["Bokermann, W. C. A. (1967) Tres novas especies de Physalaemus do sudeste brasileiro (Amphibia, Leptodactylidae). Revista Brasileira de Biologia, 27 (2), 135 - 143."]}

Physalaemus carrizorum Cardozo & Pereyra, 2018 We found a single call type for the species, referred to as call A. The call is composed of a single harmonic note, with general downward FM, but with an up-downward FM segment in the first fourth of the call. Call A (Fig. 49 A–D and 42H). We examined one recording, a total of four minutes, with 44 calls from three males. Only some of these calls were measured (see Table 2). Call duration varies from 2.360 to 4.118 s. The envelope of the call is variable. In most calls, rise and fall are similar in duration and shape (exponential) and the sustain is long. Some sustains are regular (Fig. 49C) and others are irregular (Fig. 49A), with short and shallow valleys. In some calls, the limits between the call rise, sustain, and call fall are not clear. Usually, the envelope is divided into two parts with different amplitude levels (Fig. 49A). The amplitude peak is usually at the end of the first seven tenths of the call duration. The envelope varies from elliptic or rectangular (Fig. 49C), to triangular (pointed left; Fig. 49A; rarely pointed right). Due to the asymmetry of some triangular envelopes, the shape resembles an arrow. More than 50 % of the call energy is concentrated in 36 % of the call duration around the amplitude peak. There is no PAM in the call. The call has a harmonic series (Fig. 42H). The fundamental frequency is ca. 460 Hz and approximately the first seven harmonics are emphasized. The wave periods are regular and harmonics are clear throughout the call. The dominant frequency varies from ca. 410 to 2630 Hz (Fig. 49B). The dominant harmonic is the first, fifth, or sixth (Fig. 42H). There is a clear shift in relative energy between the bands; the dominant frequency gets higher toward the end of the call, starting at the first harmonic, moving to the fifth, and ending at the sixth (Fig. 42H, 49B). Most of the call energy is between 350 and 3500 Hz (eight to nine harmonics). The call has a general downward FM (Fig. 49B, D). Additionally, calls have an up-downward FM in the first fourth of the call duration, leading to slightly arc-shaped bands in this part of the call (Fig. 49B, D), and a short downward FM at the end (Fig. 49B). The general downward FM and the initial up-downward FM result in S-shaped harmonics when considering the entire call. Some calls have a slight PFM.

Physalaemus ephippifer (Steindachner, 1864) We found a single call type for the species, referred to as call A. The call is composed of a single harmonic note with a triangular envelope that resembles an arrow-like shape. There is usually a slight PAM (without silence intervals) in the final three fourths of the call duration. The call has a general downward FM, with an up-downward FM segment in the first third or first half of the call duration. Subharmonics are always present in the first half of the call. Call A (Fig. 34 A–F and 33B). We examined two recordings, a total of one minute, with ca. 130 calls from six males. Only some of these calls were measured (see Table 2). Call duration varies from 0.466 to 0.523 s. The call rise and fall are similar in duration and shape (exponential). The sustain is composed of a long and deep valley (i.e., with a concave shape; Fig. 34A, C). The envelope varies from elliptic to triangular (pointed right). Due to the concave shape of the sustain, the triangular shape of some calls resembles an arrow (Fig. 34A). The amplitude peak is at the end of the first fourth of the call duration. More than 50 % of the call energy is concentrated in 38 % of the call duration around the amplitude peak. Some calls have an intermediate PAM only in the final three fourths of the call duration (there is no silence interval between amplitude peaks; Fig. 34A). The rate of the PAM is ca. 26 Hz, forming ca. eight cycles throughout part of the call where the PAM is present. The cycle rise and fall are similar, with amplitude peak at the middle of the cycle. The call has a harmonic series (Fig. 33B). The fundamental frequency is ca. 590 Hz and approximately the first eight harmonics are emphasized. The wave periods are regular and harmonics are clear throughout the call. Subharmonics (f 0 1/2) are present in ca. the first third or half of all calls examined (Fig. 34B, E, F). The dominant frequency varies from ca. 820 to 2630 Hz. The dominant harmonics are the first, third, fourth, fifth or sixth (usually the first or sixth; Fig. 33B, 34B). At the beginning of the call the subharmonic 1.5 is the dominant band (Fig. 34B, F). There is a clear shift in the relative energy among the bands in the second half of the call; the dominant frequency gets higher toward the end of the call, starting at the first harmonic, moving to the fifth, and ending at the sixth; thenceforth, it dominant frequency gets lower, moving to the fourth or third harmonic (Fig. 33B, 34B). Most of the call energy is between 550 and 2750 Hz (three to five harmonics). The call has a general downward FM (Fig. 34B, E). Additionally, calls have an up-downward FM in the first third or half of the call duration, yielding arc-shaped bands in this part of the call and a short downward FM at the end (Fig. 34B, E). The general downward FM and the initial up-downward FM result in S-shaped harmonics when considering the entire call. Calls have a PFM throughout the second half of the call, which is inversely proportional and synchronic to the PAM (Fig. 34A, B)., Published as part of Hepp, Fábio & Pombal, José P., 2020, Review of bioacoustical traits in the genus Physalaemus Fitzinger, 1826 (Anura: Leptodactylidae: Leiuperinae), pp. 1-106 in Zootaxa 4725 (1) on page 66, DOI: 10.11646/zootaxa.4725.1.1, http://zenodo.org/record/3612996

Physalaemus albifrons (Spix, 1824) We found a single call type for the species, referred to as call A. The call is composed of a short single harmonic note with an elliptic envelope. It has a general downward FM throughout the call but with an up-downward FM segment in the first three fourths of the call duration. Call A (Fig. 28 A–F and 24E). We examined two recordings, a total of two minutes, with ca. 90 calls from two males. Only some of these calls were measured (see Table 2). Call duration varies from 0.263 to 0.297 s. The call rise and fall durations are similar to each other in duration and shape (linear to exponential). There is a sustain, usually with a concave shape, yielding two amplitude peaks around the middle of the call (Fig. 28A). The amplitude peak is at around the middle of the call duration. The envelope is elliptic (Fig. 28A, C). More than 50 % of the call energy is concentrated in 24 % of the call duration around the amplitude peak. There is no PAM in the call. The call has a harmonic series (Fig. 24E). The fundamental frequency is ca. 480 Hz and approximately the first seven harmonics are emphasized. The wave periods are regular and harmonics are clear throughout the call. The dominant frequency varies from ca. 2440 to 2670 Hz (Fig. 28B). The dominant harmonic varies from the second to the ninth, but it is usually the fifth. There is a clear shift in the relative energy among the bands; the dominant frequency gets higher toward the end of the call, starting at the second harmonic and ending at the eighth or ninth (Fig. 28B, E, F; 24E). Thenceforth, it gets lower, ending in the seventh or sixth harmonic (Fig. 28B, E, F; 24E). Most of the call energy is between 1100 and 3700 Hz (six to nine harmonics). The call has a general downward FM (Fig. 28B, E, F). Additionally, the calls have an up-downward FM in the first three fourths of the call duration, yielding arc-shaped bands in this part of the call and a short downward FM at the end (Fig. 28B, E, F). The general downward FM and the initial up-downward FM result in S-shaped harmonics when considering the entire call. There is no PFM., Published as part of Hepp, Fábio & Pombal, José P., 2020, Review of bioacoustical traits in the genus Physalaemus Fitzinger, 1826 (Anura: Leptodactylidae: Leiuperinae), pp. 1-106 in Zootaxa 4725 (1) on page 60, DOI: 10.11646/zootaxa.4725.1.1, http://zenodo.org/record/3612996

Physalaemus erikae Cruz & Pimenta, 2004 We found a single call type for the species, referred to as call A. The call is composed of a single harmonic note. It has a general downward FM throughout the call but with an up-downward FM segment in the first sixth of the call duration. Call A (Fig. 29 A–H and 24F). We examined two recordings, a total of four minutes, with 37 calls from four males. Only some of these calls were measured (see Table 2). Call duration varies from 0.478 to 0.566 s. The limits between the call rise, sustain, and call fall are not clear (mainly in calls with elliptic envelope, see Fig. 29C, D). When perceptible, the call rise and fall can be similar in duration, or fall shorter than rise. Usually, call rise has an exponential shape, whereas call fall has a logarithmic shape. When present, the sustain is irregular, usually with short and very shallow amplitude valleys (Fig. 29A, E). The amplitude peak is at around the middle of the call duration. The envelope of the call varies from elliptic (Fig. 29C, D, E) to slightly rectangular (when flat sustains are present; Fig. 29A). More than 50 % of the call energy is concentrated in 30 % of the call duration around the amplitude peak. There is no PAM in the call. The call has a harmonic series (Fig. 24F). The fundamental frequency is ca. 420 Hz and approximately the first eight harmonics are emphasized. The wave periods are regular and harmonics are clear throughout the call. The dominant frequency varies from ca. 2840 to 2890 Hz (Fig. 29B). The dominant harmonic varies from the first to the seventh. There is a clear shift in the relative energy among bands; the dominant frequency gets higher toward the end of the call, starting at the first harmonic and ending at the seventh (Fig. 24F, 29B, F, G, H). Most of the call energy is between 650 and 3450 Hz (five to six harmonics). The call has a general downward FM (Fig. 29B, F, G, H). Additionally, the calls have an up-downward FM in the first sixth of the call duration, yielding arc-shaped bands in this part of the call, and a short downward FM at the end (Fig. 29B, F, G, H). The general downward FM and the initial up-downward FM result in S-shaped harmonics when considering the entire call. There is no PFM., Published as part of Hepp, Fábio & Pombal, José P., 2020, Review of bioacoustical traits in the genus Physalaemus Fitzinger, 1826 (Anura: Leptodactylidae: Leiuperinae), pp. 1-106 in Zootaxa 4725 (1) on page 61, DOI: 10.11646/zootaxa.4725.1.1, http://zenodo.org/record/3612996, {"references":["Pimenta, B. V. S. & Cruz, C. A. G. (2004) The tadpole and advertisement call of Physalaemus aguirrei Bokermann, 1966 (Amphibia, Anura, Leptodactylidae). Amphibia-Reptilia, 25, 197 - 204. https: // doi. org / 10.1163 / 1568538041231201"]}

Physalaemus moreirae (Miranda-Ribeiro, 1937) We found two different calls, referred to as call A and B. Calls A and B are composed of a single harmonic note each. Call A is composed of pulses whereas Call B is not. Moreover, call B has irregular FM segments (vs. absent in call A). Call A (Fig. 21 A–J and 13H). We examined three recordings, a total of one minute, with 68 calls from five males. Only some of these calls were measured (see Table 2). Call duration varies from 0.499 to 0.567 s. In most calls, the limits between call rise, sustain, and fall are not clear (for example in calls with elliptic envelope; Fig. 21E). When perceptible, the call rise and fall are similar in duration and shape, both with a linear shape, and there is a long sustain. This segment can be regular and flat (Fig. 21A) or quite irregular with AM segments within it (Fig. 21C). The amplitude peak is usually at around the middle of the call duration. The envelope varies from elliptic (Fig. 21E) to triangular (pointed left; Fig. 21C). More than 50 % of the energy is concentrated in 35 % of the call duration around the amplitude peak. The call has a strong PAM (with silence intervals present between pulses; Fig 21A, C, E). The rate of the PAM is ca. 42 Hz, forming ca. 23 pulses throughout the call. Most pulses have an elliptic envelope with the amplitude peak around the middle of the pulse. The very first pulses have rise longer than fall, amplitude peak at the end of pulse and the very last pulses have the opposite shape (amplitude peak at the beginning of the pulse; Fig. 21D, F). The last pulse is the longest (Fig. 21 A–J). Short silence intervals are present between the pulses, but can be absent between the first and last pulses (pulses are juxtaposed to neighboring pulses; Fig. 21A, C, D, E, F). The call has a harmonic series (Fig. 13H). The fundamental frequency is ca. 230 Hz and this band can be present with low energy or absent in the audiospectrograms. Usually, the wave periods are regular and harmonics are clear throughout the call. However, the short duration of the pulses (except the last) make the bands broad with narrow intervals. Where two pulses are juxtaposed, the wave periods are less regular, and the harmonics are less clear with deterministic chaos (Fig. 21B, G, H, I, J). The dominant frequency varies from ca. 770 to 1250 Hz (Fig. 21B). The dominant harmonic varies from the third to the eighth, but it is usually the third or fourth. There is no clear shift in the relative energy among the bands throughout the call. Most of the energy is concentrated between 700 and 1300 Hz (three harmonics). A few calls have a slight upward general FM (Fig. 21H, I). Usually, there is no clear general FM (Fig. 21B). Calls have a short downward FM at the end (Fig. 21H, J) and PFM throughout the call without clear relationship with the synchronic pulse-PAM (Fig. 21E, F, I, J). Call B (Fig. 21 K–L and 16E). We examined one recording, a total of one minute, with one call from one male. This call was also measured (see Table 2). A single call was recorded. Call duration is 0.254 s. The call rise and fall are similar in duration and shape (exponential-shaped). There is a long sustain, which is regular until the amplitude peak at the end of the call, around nine tenths of the call duration. The envelope is triangular (pointed left; Fig. 21K). More than 50 % of the energy is concentrated in 54 % of the call duration around the amplitude peak. This call has no PAM. The call has a harmonic series (Fig. 16E). The fundamental frequency is at ca. 280 Hz and this band is generally absent in the audiospectrogram. The wave periods are regular and harmonics are clear throughout the call. However, the wave periods are less regular at the middle of the call, with biphonation regime (Fig. 21L). The dominant frequency is ca. 860 Hz (Fig. 21L). The dominant harmonic varies from the second to the fifth, but it is usually the third or fourth. There is no clear shift in the relative energy among the bands throughout the call. Most of the energy is concentrated between 800 and 1400 Hz (often, three harmonics). The long downward FM segment at the beginning makes the general FM of the call downward. There is a slight PFM throughout the call. Additionally, the call has parts with irregular up and downward FM (Fig. 21L)., Published as part of Hepp, Fábio & Pombal, José P., 2020, Review of bioacoustical traits in the genus Physalaemus Fitzinger, 1826 (Anura: Leptodactylidae: Leiuperinae), pp. 1-106 in Zootaxa 4725 (1) on pages 52-54, DOI: 10.11646/zootaxa.4725.1.1, http://zenodo.org/record/3612996

Physalaemus cuqui Lobo, 1993 We found a single call type for the species, referred to as call A. The call has a single harmonic note with a slight PAM without silence intervals. It has a gradual downward FM throughout the call. Call A (Fig. 26 A–B and 24C). We examined two recordings, a total of two minutes, with 47 calls from six males. Only some of these calls were measured (see Table 2). Call duration varies from 1.215 to 1.500 s. The limits between the call rise, sustain, and call fall are not clear (see elliptic envelope in Fig. 26A); the envelope is linear- or exponential-shaped until the amplitude peak and logarithmic-shaped from the peak to the end of the call (Fig. 26A). The amplitude peak is at around the end of the first two thirds of the call duration. The envelope is elliptic (Fig. 26A). More than 50 % of the call energy is concentrated in 26 % of the call duration around the amplitude peak. The call has a slight PAM (there is no silence interval between amplitude peaks; Fig. 26A). The rate of the PAM is ca. 21 Hz, yielding ca. 29 cycles throughout the call. The cycle rise and fall are similar, with amplitude peak at the middle of the cycle. The call has a harmonic series (Fig. 24C). The fundamental frequency is ca. 510 Hz and approximately the first seven harmonics are emphasized. The wave periods are regular and harmonics are clear throughout the call. The dominant frequency is ca. 2850 Hz (Fig. 26B). The dominant harmonic varies from the first to the sixth, but it is usually the sixth. There is a clear shift in relative energy among the bands; the dominant frequency gets higher toward the end of the call, starting at first harmonic and ending in the sixth one (Fig. 26B). Most of the call energy is between 500 and 3000 Hz (five to six harmonics). The call has a general downward FM. Additionally the calls have a subtle up-downward FM at the beginning, yielding a arc-shaped bands in this part of the call in audiospectrograms, and a short downward FM at the end (Fig. 26B). The general downward FM and the initial up-downward FM result in S-shaped harmonics when considering the entire call (Fig. 26B). The call also has a PFM, which is inversely proportional and synchronic to the PAM (26A–B)., Published as part of Hepp, Fábio & Pombal, José P., 2020, Review of bioacoustical traits in the genus Physalaemus Fitzinger, 1826 (Anura: Leptodactylidae: Leiuperinae), pp. 1-106 in Zootaxa 4725 (1) on page 58, DOI: 10.11646/zootaxa.4725.1.1, http://zenodo.org/record/3612996

Physalaemus maculiventris (A. Lutz, 1925) We found two different calls, referred to as call A and B. Calls B were common in recordings in which several males were more active and calling close to each other. Calls B are commonly observed after overlapping periods of A calls. Call B resembles a long call A with higher fundamental frequency, pulse-PAM rate, and stronger PFM. Call A (Fig. 5 A–D and 4B). We examined two recordings, a total of six minutes, with ca. 400 calls from four males. Only some of these calls were measured (see Table 2). Call duration varies from 0.172 to 0.260 s. The rise of the call is longer than the fall; the amplitude peak is approximately at the end of the first three fourths of the call duration. Since both rise and fall are relatively similar in slope and not too different in duration, the envelope of the call is fairly elliptic (Fig. 5A, C). More than 50 % of the call energy is concentrated in 24 % of the call duration around the amplitude peak. The call has a subtle PAM (there is no silence interval between peaks; Fig. 5C). The rate of the PAM is ca. 39 Hz, forming ca. six amplitude peaks throughout the call. The call is composed of usually clear harmonics (Fig. 4B), however, eventual decreases of the wave periodicity make the harmonics less clear. The fundamental frequency is approximately 220 Hz and it can be present with low energy or absent in the audiospectrograms. The dominant frequency varies from ca. 820 to 1510 Hz (Fig. 5B, D). The dominant harmonic varies from the third to the sixth, but it is usually the fifth. There is no clear shift in the relative energy among the bands throughout the call. Most of the call energy is between 700 and 1500 Hz (often comprising five harmonics). The frequency bands have a general upward FM throughout the call with a rapid up-downward FM at the beginning forming arc-shaped bands in this part of the call and a short downward-FM segment at the end (Fig. 5B). There are irregular PFM segments throughout the entire call; these segments are usually synchronic and directly proportional to the PAM (Fig. 5D). ...Continued on the next page ...Continued on the next page ...Continued on the next page ...Continued on the next page ...Continued on the next page ...Continued on the next page ...Continued on the next page ...Continued on the next page ...Continued on the next page ...Continued on the next page ...Continued on the next page ...Continued on the next page Call B (Fig. 5 E–F and 6A). We examined one recording, a total of one minute, with six calls from two males. Only some of these calls were measured (see Table 2). Call duration varies from 0.375 to 0.675 s. The call rise is longer than the fall with a long regular sustain or shallow valley separating them; the shallow valley yields two amplitude peaks at the beginning and end of the call. The amplitude peak is at around the end of the first two thirds of the call duration (i.e., second amplitude peak). Depending on the slope of the sustain and the difference between the peaks, the envelope of the call can vary from rectangular to triangular (pointed left; Fig. 5E). More than 50 % of the call energy is concentrated in 34 % of the call duration around the amplitude peak. The call has a slight PAM (there is no silence interval between the peaks; Fig. 5E). The rate of the PAM is ca. 32 Hz, forming ca. 13 amplitude peaks throughout the call. The call is composed of harmonics (Fig. 6A). Usually the harmonics are clear, however, eventual decreases of the wave periodicity make the harmonics less clear. Audiospectrograms with relatively broad filter bandwidths (e.g., above 100 Hz) can show wave peaks, of some parts of the call with low fundamental frequencies (minimum 123 Hz; see Table 2), as broadband pulses (instantaneously high sound-pressure effect; see Littlejohn 2001). The fundamental frequency is usually ca. 240 Hz and it can be present with low energy or absent in the audiospectrograms. The dominant frequency varies from ca. 1030 to 1310 Hz (Fig. 5F). The dominant harmonic varies from the third to the eighth, but it is usually the fourth or the fifth harmonic. There is no clear shift in the relative energy among the bands throughout the call. Most of the call energy is between 800 and 1600 Hz (ca. three harmonics). The frequency bands have a general upward FM throughout the call with a up-downward FM segment at the beginning forming arc-shaped bands in this part of the call and a short downward FM segment at the end (Fig. 5F). There are irregular PFM segments throughout the entire call; these segments are usually synchronic and directly proportional to the PAM (Fig. 5E, F)., Published as part of Hepp, Fábio & Pombal, José P., 2020, Review of bioacoustical traits in the genus Physalaemus Fitzinger, 1826 (Anura: Leptodactylidae: Leiuperinae), pp. 1-106 in Zootaxa 4725 (1) on pages 15-32, DOI: 10.11646/zootaxa.4725.1.1, http://zenodo.org/record/3612996, {"references":["Lourenco, L. B., Targueta, C. P., Baldo, D., Nascimento, J., Garcia, P. C. A., Andrade, G. V., Haddad, C. F. B. & Recco-Pimentel, S. M. (2015) Phylogeny of frogs from the genus Physalaemus (Anura, Leptodactylidae) inferred from mitochondrial and nuclear gene sequences. Molecular Phylogenetics and Evolution, 92, 204 - 216. https: // doi. org / 10.1016 / j. ympev. 2015.06.011","Littlejohn, M. J. (2001) Pattern of differentiation in temporal properties of Acoustic signals of anurans. In: Michael, J. R. (Ed.), Anuran Communication. Smithsonian Institution Press, Washington, pp. 102 - 120."]}

Physalaemus angrensis Weber, Gonzaga & Carvalho-e-Silva, 2006 We found two different calls, referred to as call A and B. Calls A and B are composed of harmonics and a single note each. Call A is composed of pulses whereas Call B has no PAM. Moreover, call B has a stronger general upward FM. Call B can have irregular FM segments and jumps of the fundamental frequency (vs. absent in call A). Call A (Fig. 19 A–H and 13F). We examined nine recordings, a total of 32 minutes, with ca. 800 calls from 19 males. Only some of these calls were measured (see Table 2). Call duration varies from 0.431 to 0.605 s. In most calls, the limits between the call rise, sustain, and fall are not clear (for example in calls with elliptic envelopes; see below; Fig. 19D). In calls where the limits are perceptible, the call rise and fall are similar in duration and shape, both have a logarithmic or linear shape, and there is a long sustain, which can have shallow valleys and short slopes (concave and convex shapes, respectively; Fig. 19A). The amplitude peak is at around the end of the first three fourths of the call duration. The envelope varies from elliptic (Fig. 19D) to rectangular (Fig. 19A) depending on how emphasized or regular is the sustain. More than 50 % of the energy is concentrated in 37 % of the call duration around the amplitude peak. This call has a strong PAM (there are silence intervals present between pulses; Fig. 19A, B, D, E, G, H). The rate of the PAM is ca. 58 Hz, yielding ca. 30 pulses throughout the call. Except for the last pulse, the pulse rise is longer than the fall and the amplitude peak is around two thirds of the pulse duration. The last pulse has the opposite envelope with amplitude peak at the beginning (Fig. 19E). The first pulses can have much lower amplitude than the others. The last pulse is the longest. There are short silence intervals between pulses, which can be absent between the first and last pulses (pulses are juxtaposed to neighboring pulses; Fig. 19A, B, D, E, G, H). Intervals are usually eightfold longer than the pulse durations. The call has a harmonic series (Fig. 13F). The fundamental frequency is at ca. 410 Hz and this band can be present with low energy or absent in the audiospectrograms. The wave periods are regular and harmonics are clear throughout the call. The dominant frequency varies from ca. 1590 to 1780 Hz (Fig. 19B). The dominant harmonic varies from the third to the fifth, but it is usually the fourth. There is no clear shift in the relative energy among the bands throughout the call. Most of the energy is concentrated between 1200 and 1900 Hz (three harmonics). The call has a general upward FM (Fig. 19B, G). Additionally, there is PFM throughout the call, which is usually directly proportional to the synchronic pulse-PAM (Fig. 19E, H). Calls are usually emitted in short sequences with up to ten calls each (Fig. 19C, F). Call B (Fig. 19 I–L and 16D). We examined three recordings, a total of 18 minutes, with ca. 150 calls from five males. Only some of these calls were measured (see Table 2). Call duration varies from 0.309 to 0.353 s. The call rise and fall are similar in duration and shape (logarithmic-shaped). There is a sustain, which has shallow valleys, usually at its beginning and end (Fig. 19I, K). The amplitude peak of the call is at around the end of the first four fifths of the call duration (Fig. 19I, K). The envelope varies from elliptic to triangular (pointed left; Fig. 19I, K). More than 50 % of the energy is concentrated in 32 % of the call duration around the amplitude peak. This call has no PAM. The call has a harmonic series (Fig. 16D). The fundamental frequency is ca. 320 Hz and this band can be present with low energy or absent in the audiospectrograms. The wave periods are regular and harmonics are clear throughout the call. The dominant frequency varies from ca. 1080 to 1310 Hz (Fig. 19J). The dominant harmonic varies from the second to the seventh harmonic, but it is usually the fourth. There is no clear shift in the relative energy among the bands throughout the call. Most of the energy is concentrated between 900 and 1400 Hz (two harmonics). The call has a general upward FM with short downward FM at the end (Fig. 19J, L). The sustain has an up-downward FM (Fig. 19J, L). There is clear PFM in some parts of the call. Additionally, several calls have parts with irregular up and downward FM, usually, inversely proportional to the AM directions (Fig. 19 I–L)., Published as part of Hepp, Fábio & Pombal, José P., 2020, Review of bioacoustical traits in the genus Physalaemus Fitzinger, 1826 (Anura: Leptodactylidae: Leiuperinae), pp. 1-106 in Zootaxa 4725 (1) on pages 50-51, DOI: 10.11646/zootaxa.4725.1.1, http://zenodo.org/record/3612996, {"references":["Weber, L. N., Gonzaga, L. P. & Carvalho-E-Silva, S. P. (2006) \" 2005 \" A new species of Physalaemus Fitzinger, 1826 from the lowland Atlantic Forest of Rio de Janeiro state, Brazil (Amphibia, Anura, Leptodactylidae). Arquivos do Museu Nacional, 63 (4), 677 - 684."]}

Physalaemus maximus Feio, Pombal & Caramaschi, 1999 We found a single call type for the species, referred to as call A. The call is composed of a single harmonic note with a subtle PAM, with no silence intervals. It has a long duration and a very low fundamental frequency with subtle PFM throughout the call. The bands have a slight general upward FM and a downward FM at the end, yielding a slight arc shape in the audiospectrogram when considering the entire call. Call A (Fig. 39 A–D and 33G). We examined seven recordings, a total of five minutes, with ca. 90 calls from 11 males. Only some of these calls were measured (see Table 2). Call duration varies from 1.887 to 2.446 s. The call rise is longer than call fall or both are similar in duration. Call rise and fall have exponential, linear or logarithmic shape. There is a long sustain. It is usually almost flat but slightly irregular (Fig. 39A). However, in some calls, the beginning of this segment has low amplitude, which gradually increases towards the end of the call (Fig 39C). The amplitude peak is usually at the end of the first two thirds the call duration. The amplitude of the call is usually at three fifths of the call duration. The envelope varies from elliptic, rectangular (Fig. 39A) to triangular (pointed left; Fig. 39C) depending on the steepness of the sustain and position of the amplitude peak of the call. More than 50 % of the call energy is concentrated in 36 % of the call duration around the amplitude peak. The call can have a slight PAM (silence intervals absent between peaks). The rate of the PAM is ca. 10 Hz, forming ca. 22 amplitude peaks throughout the call. The call has a harmonic series (Fig. 33G). The fundamental frequency is ca. 170 Hz. This band and the next harmonic are absent in audiospectrograms. There are usually ca. seven emphasized harmonics. The wave periods are regular and harmonics are clear throughout the call. The dominant frequency varies from ca. 1000 to 1030 Hz (Fig. 39B). The dominant harmonic varies from the third to the ninth, but it is usually the sixth (Fig. 33G). There is no clear shift in the relative energy between the bands throughout the call. Most of the call energy is between 850 and 1550 Hz (five harmonics). The call has a general FM slightly upward and a short downward FM at the end, yielding a slight arc shape in the audiospectrogram when considering the entire call (Fig. 39B, D). Additionally, there can be a slight PFM throughout the call, which is usually independent of PAM or can be directly proportional and synchronic to some parts of the PAM., Published as part of Hepp, Fábio & Pombal, José P., 2020, Review of bioacoustical traits in the genus Physalaemus Fitzinger, 1826 (Anura: Leptodactylidae: Leiuperinae), pp. 1-106 in Zootaxa 4725 (1) on page 71, DOI: 10.11646/zootaxa.4725.1.1, http://zenodo.org/record/3612996

Physalaemus erythros Caramaschi, Feio & Guimarães, 2003 We found two different calls, referred to as call A and B. B calls were common in recordings in which several males were active and calling close to each other. B calls were commonly observed after overlapping periods of calls A. Call B differs from call A by its longer duration, higher fundamental frequency, presence of pulse-PAM and PFM. Additionally, the envelope of A calls is elliptic whereas that of the B calls is triangular or rectangular. Call A (Fig. 7 A–D and 4D). We examined 21 recordings, a total of 56 minutes, with ca. 4900 calls from 15 males. Only some of these calls were measured (see Table 2). Call duration varies from 0.041 to 0.077 s. The call rise and fall are similar in duration, rendering an elliptic call envelope; the call fall is gradual, whereas the call rise is steeper. The amplitude peak is at around the middle of the call duration (Fig. 7A, C). More than 50 % of the call energy is concentrated in 39 % of the call duration around the amplitude peak. The call has no PAM. The call is composed of harmonics (Fig. 4D). The harmonics are very close to each other and hardly distinguished due to the low fundamental frequency and the lack of the wave periodicity throughout the call. The fundamental frequency is approximately 250 Hz and this band can be present with low energy or absent in the audiospectrograms. The dominant frequency varies from ca. 1020 to 1160 Hz (Fig. 7B, D). The dominant harmonic varies from the second to the tenth, but it is usually the third. There is no clear shift in the relative energy among the bands throughout the call. Most of the call energy is between 300 and 2000 Hz (ca. seven harmonics). There is no clear general FM in the call but there are subtle irregular FM segments throughout the entire call. Call B (Fig. 7 E–J and 6B). We examined eight recordings, a total of 26 minutes, with 93 calls from eight males. Only some of these calls were measured (see Table 2). Call duration varies from 0.200 to 0.875 s. The call rise is abrupt and a little shorter than the fall, which is also very short but gradual. The call has a long sustain. The amplitude is usually regular throughout the call (Fig. 7H). However, in some calls, the amplitude increases gradually toward the end of the call (Fig. 7E, G). The amplitude peak is at the very end of the call duration (Fig. 7E, G, H). Depending on the slope of the sustain and the difference between the amplitude peaks, the envelope of the call can vary from rectangular (Fig. 7H) to triangular (pointed left; Fig. 7E, G). More than 50 % of the call energy is concentrated in 29 % of the call duration around the amplitude peak. The call has a weak to intermediate PAM (there is no silence interval between the peaks; Fig. 7E, G, H). The rate of the PAM is ca. 27 Hz, forming ca. 13 amplitude peaks throughout the call. The envelope of this PAM cycles is variable but the amplitude peak is at the middle of the cycle. The call is composed of harmonics (Fig. 6B). Usually the harmonics are clear, however, eventual decrease in the wave periodicity makes the harmonics less clear with deterministic chaos regimes. Audiospectrograms with relatively broad filter bandwidths (e.g., above 100 Hz) can show wave peaks in some parts of the call with low fundamental frequencies (minimum 107 Hz; see Table 2), as broadband pulses (instantaneously high sound-pressure effect; see Littlejohn 2001). The fundamental frequency is around 250 Hz and it is usually absent in the audiospectrograms. The dominant frequency varies from ca. 840 to 1780 Hz (Fig. 7F, I, J). The dominant harmonic varies from the third to the ninth, but it is usually the third and fourth harmonic. There is no clear shift in the relative energy among the bands throughout the call. Most of the call energy is between 800 and 1600 Hz (three to four harmonics). The frequency bands have a general upward FM throughout the call and a short downward FM at the end (Fig. 7F, I, J). There are irregular PFM segments throughout the entire call; these segments are usually synchronic and directly proportional to the PAM (Fig. 7 E–J)., Published as part of Hepp, Fábio & Pombal, José P., 2020, Review of bioacoustical traits in the genus Physalaemus Fitzinger, 1826 (Anura: Leptodactylidae: Leiuperinae), pp. 1-106 in Zootaxa 4725 (1) on pages 34-35, DOI: 10.11646/zootaxa.4725.1.1, http://zenodo.org/record/3612996, {"references":["Caramaschi, U., Feio, R. N. & Guimaraes, A. S. N. (2003) A new, brightly colored species of Physalaemus (Anura: Leptodactylidae) from Minas Gerais, southeastern Brazil. Herpetologica, 59, 519 - 524. https: // doi. org / 10.1655 / 02 - 102","Littlejohn, M. J. (2001) Pattern of differentiation in temporal properties of Acoustic signals of anurans. In: Michael, J. R. (Ed.), Anuran Communication. Smithsonian Institution Press, Washington, pp. 102 - 120."]}

Physalaemus nattereri (Steindachner, 1863) We found a single call type for the species, referred to as call A. The call is composed of a single harmonic note usually emitted in sequences of three or four calls (Fig. 3A and B), but longer sequences, with dozens of calls, can be sporadically emitted. Call A (Fig. 3 A–F and 4A). We examined 15 recordings, a total of 21 minutes, with ca. 5500 calls from 28 males. Only some of these calls were measured (see Table 2). Call duration varies from 0.057 to 0.074 s. The abrupt call rise is much shorter than the call fall, which is more gradual; the amplitude peak is within the first third of the call’s duration (usually at the very beginning of the call). The envelope of the call resembles a triangle pointed right. More than 50 % of the call energy is concentrated at the first 26 % of the call duration (Fig. 3C, E). The call has no PAM. The call is composed of clear harmonics (Fig. 3D, F, 4A). The fundamental frequency is around 330 Hz and it is usually present with low energy or absent in the audiospectrograms. The dominant frequency of the calls varies from 703 to 1031 Hz. Within each call, the dominant harmonic varies from the first to third, but it is usually the second or third. There is no clear shift in the relative energy among the bands throughout the call. Most of the call energy is between 700 and 1050 Hz (often, two harmonics). Frequency modulations are very subtle in this species. Nonetheless, the frequency bands have a general downward FM with a slight rapid up-downward FM at the beginning and a short down or upward FM at the end. There is no PFM (Fig. 3D, F)., Published as part of Hepp, Fábio & Pombal, José P., 2020, Review of bioacoustical traits in the genus Physalaemus Fitzinger, 1826 (Anura: Leptodactylidae: Leiuperinae), pp. 1-106 in Zootaxa 4725 (1) on page 15, DOI: 10.11646/zootaxa.4725.1.1, http://zenodo.org/record/3612996

Physalaemus atim Brasileiro & Haddad, 2015 We found a single call type for the species, referred to as call A. The call has a single harmonic note with a gradual downward FM throughout the call. Call A (Fig. 27 A–D and 24D). We examined one recording, a total of 11 seconds, with 11 calls from four males. Most of these calls were measured (see Table 2). Call duration varies from 0.779 to 0.995 s. The limits between the call rise, sustain and call fall are not clear (Fig. 27A). In calls where they are perceptible, the call rise and fall can be similar in duration with variable shape (linear, exponential, or logarithmic) or fall is shorter than rise. The sustain is irregular with short amplitude valleys (Fig. 27A). The amplitude peak is at around the end of the first three fifths of the call duration. The envelope is elliptic (Fig. 27A), rectangular (Fig. 27C) or triangular (pointed left). More than 50 % of the call energy is concentrated in 33 % of the call duration around the amplitude peak. The call has an irregular slight PAM (there is no silence interval between amplitude peaks). The rate of the PAM is ca. 45 Hz, forming ca. 24 peaks throughout the call. The cycle rise and fall are similar, with amplitude peak at the middle of the cycle. The call has a harmonic series (Fig. 24D). The fundamental frequency is ca. 430 Hz and approximately the first eight harmonics are emphasized. The wave periods are regular and harmonics are clear throughout the call. The dominant frequency varies from ca. 1980 to 2330 Hz (Fig. 27B). The dominant harmonic varies from the first to the seventh, but it is usually the sixth. There is a clear shift in relative energy among the bands; the dominant frequency gets higher toward the end of the call, starting at the first harmonic and ending at the sixth (Fig. 27D). Most of the call energy is between 700 and 2700 Hz (five to six harmonics). The call has a general downward FM (Fig. 27B, D). Additionally, the calls have a subtle up-downward FM at the beginning, yielding arc-shaped bands in this part of the call (Fig. 27D), and a short downward FM at the end (Fig. 27B, D). The general downward FM and the initial up-downward FM result in S-shaped harmonics when considering the entire call. There is no clear PFM., Published as part of Hepp, Fábio & Pombal, José P., 2020, Review of bioacoustical traits in the genus Physalaemus Fitzinger, 1826 (Anura: Leptodactylidae: Leiuperinae), pp. 1-106 in Zootaxa 4725 (1) on page 59, DOI: 10.11646/zootaxa.4725.1.1, http://zenodo.org/record/3612996, {"references":["Brasileiro, C. A. & Haddad, C. F. B. (2015) A New Species of Physalaemus from Central Brazil (Anura: Leptodactylidae). Herpetologica, 71 (4), 280 - 288. https: // doi. org / 10.1655 / HERPETOLOGICA-D- 13 - 00085"]}

Physalaemus crombiei Heyer & Wolf, 1989 We found two different calls, referred to as call A and B. B calls were observed in recordings in which males emitted A calls with very long durations. Calls A and B are composed of harmonics and a single note each. A calls have pulses separated by silence intervals whereas B calls have not. Moreover, B calls have a general upward FM and FM segments (periodic or not) stronger than those of A calls. Call A (Fig. 15 A–J and 13C). We examined nine recordings, a total of 20 minutes, with ca. 900 calls from 18 males. Only some of these calls were measured (see Table 2). Call duration varies from 0.319 to 0.774 s. The call rise and fall durations are usually similar to each other and they can be gradual or abrupt, linear or logarithmic; there is a long sustain (Fig. 15A, C, D, E). This segment can be slightly concave or convex (Fig. 15D, E, respectively). The amplitude peak is often at around (usually just after) the middle of the call duration. Since both rise and fall are similar in slope and duration, the envelope of the call is fairly elliptic (Fig. 15E) but can be rectangular (Fig. 15C, D) or triangular (pointed left; Fig. 15A) depending on the shape of sustained segment and the position of the amplitude peak of the call. More than 50 % of the call energy is concentrated in 43 % of the call duration around the amplitude peak. The call has a strong PAM (with silence intervals present between pulses; Fig. 15 A–J). The rate of this PAM is ca. 25 Hz, forming ca. nine pulses throughout the call. Except for the last pulse, the rise of the pulses is longer than the fall and the amplitude peak is at around the end of the first two thirds of the pulse duration (Fig. 15F). The last pulse has the inverse envelope with amplitude peak at its outset (Fig. 15F). In some calls, the last pulse is notably longer than the others (Fig. 15A, B, C, E, F, G, I, J). Silence intervals are present between pulses, ca. tenfold shorter than pulse duration (Fig. 15 A–J). The call has a harmonic series (Fig. 13C). The fundamental frequency is ca. 370 Hz and this band can be present with low energy or absent in the audiospectrograms. The wave periods are very regular and the harmonics are clear throughout the call. Jumps of the fundamental frequency can be present between the first pulses. The dominant frequency varies from ca. 1010 to 1380 Hz (Fig. 15B). The dominant harmonic varies from the third to the fourth, but it is usually the third. There is no clear shift in the relative energy between the bands throughout the call. Most of the call energy is between 900 and 1300 Hz (two harmonics). The frequency bands have a general upward FM throughout the call and short downward FM at the end (Fig. 15B, G, H, I). Additionally, there is PFM throughout the call, which is directly proportional to the synchronic pulse-PAM (Fig. 15B, G, H, I, J). Call B (Fig. 15 K–N and 16A). We examined one recording, a total of five minutes, with two calls from one male. Most of these calls were measured (see Table 2). Call duration varies from 0.709 to 0.945 s. The envelope of the call is variable; call rise and fall are short. There can be more than one sustain, with different amplitudes (Fig. 15K, M). Usually, the first has lower amplitude (Fig. 15K). The amplitude peak is at around the middle or beginning of the call duration. The envelope can be classified as elliptic, triangular (pointed right; Fig. 15M) or rectangular (Fig. 15K). More than 50 % of the call energy is concentrated in 30 % of the call duration around the amplitude peak. One call clearly shows a section with a slight PAM (there is no silence interval between the amplitude peaks; Fig. 15K). The rate of this PAM is ca. 19 Hz, forming ca. seven emphasized peaks at the middle of the call duration. The call has a harmonic series (Fig. 16A). The fundamental frequency is ca. 340 Hz and this band can be present with low energy or absent in the audiospectrograms. One call shows a sudden jump of the fundamental frequency at the end of the call. The wave periods are regular and then the harmonics are clear throughout the call. The dominant frequency varies from ca. 1020 to 1160 Hz (Fig. 15L). The dominant harmonic is the third. There is no clear shift in the relative energy between the bands throughout the call. Most of the call energy is between 600 and 2000 Hz (ca. five harmonics). The frequency bands have a general upward FM throughout the call and short downward FM at the end (Fig. 15L, N). Additionally, there is PFM throughout the call, which is directly proportional to the synchronic pulse-PAM where it is present (15K–N)., Published as part of Hepp, Fábio & Pombal, José P., 2020, Review of bioacoustical traits in the genus Physalaemus Fitzinger, 1826 (Anura: Leptodactylidae: Leiuperinae), pp. 1-106 in Zootaxa 4725 (1) on pages 44-46, DOI: 10.11646/zootaxa.4725.1.1, http://zenodo.org/record/3612996, {"references":["Heyer, W. R. & Wolf, A. J. (1989) Physalaemus crombiei (Amphibia: Leptodactylidae), a new frog species from Espirito Santo, Brazil with comments on the P. signifer group. Proceedings of the Biological Society of Washington, 102 (2), 500 - 506."]}

Physalaemus camacan Pimenta, Cruz & Silvano, 2005 We found a single call type for the species, referred to as call A. The call has a single harmonic note with a slight PAM. It is spectrally polymorphic with clear harmonics, sidebands, and deterministic-chaos regime. Call A (Fig. 9 A–D and 4E–F). We examined two recordings, a total of five minutes, with ca. 130 calls from two males. Only some of these calls were measured (see Table 2). Call duration varies from 0.676 to 0.980 s. The call rise is gradual and longer than the call fall, which is more abrupt. There is a long sustain in the call. Usually the amplitude of the call is regular throughout the call (Fig. 9A). However, in some calls, the amplitude increases gradually toward the amplitude peak at the end of the call (Fig. 9C). The amplitude peak is at around four fifths of the call duration. Depending on the slope of the sustain and the difference between the amplitude peaks the envelope of the call can vary from rectangular (Fig. 9A) to triangular (pointed left; Fig. 9C). More than 50 % of the call energy is concentrated in 38 % of the call duration around the amplitude peak. The call has a slight PAM (with no silence interval between peaks; Fig. 9A, C). The rate of the PAM is ca. 13 Hz, forming ca. 10 amplitude peaks throughout the call. The calls can have two different spectral patterns (Fig. 4E, F). The bands of one of these patterns (Fig. 9B) are multiple of each other and were considered harmonics. The fundamental frequency of this series is ca. 400 Hz (Fig. 9B). In the other spectral pattern (Fig. 9D), there is a series of bands with fundamental frequency of ca. 100 Hz, which varies continuously and the bands are not integral multiple of each other. The bands of this 100 Hz series seem to be sidebands (i.e., 100 Hz wave as the modulating signal) with the 410 Hz series as the carrier signal (Fig. 9D). In most calls, the sidebands are the only bands noticeable. In these calls, the bands are not very clear since there is considerably deterministic chaos (Fig. 9D) due to the irregularity of the wave periods of the 100 Hz signal. In the calls where the 400 Hz series are evident, the harmonics are clear due to the higher fundamental frequency and the more regularity (periodicity) of the wave periods. The dominant frequency varies from ca. 1380 to 1660 Hz (Fig. 9B). Considering the 400 Hz series, the dominant harmonic varies from the second to the sixth, but it is usually the fourth. There is no clear shift in the relative energy between the bands throughout the call. Most of the call energy is between 1100 and 1800 Hz. This bandwidth corresponds to two harmonics of the 400-Hz series. The frequency bands have a general upward FM throughout the call with a short downward FM at the end (Fig. 9B). There is a PFM in the parts of the call where the bands are clear (Fig. 9B). This PFM is synchronic and directly proportional to the PAM (Fig. 9A, B)., Published as part of Hepp, Fábio & Pombal, José P., 2020, Review of bioacoustical traits in the genus Physalaemus Fitzinger, 1826 (Anura: Leptodactylidae: Leiuperinae), pp. 1-106 in Zootaxa 4725 (1) on page 37, DOI: 10.11646/zootaxa.4725.1.1, http://zenodo.org/record/3612996, {"references":["Pimenta, B. V. S., Cruz, C. A. G. & Silvano, D. L. (2005) A new species of the genus Physalaemus Fitzinger, 1826 (Anura, Leptodactylidae) from the Atlantic Rain Forest of southern Bahia, Brazil. Amphibia-Reptilia, 26, 201 - 210. https: // doi. org / 10.1163 / 1568538054253483"]}

Given the importance of acoustic communication in intraspecific recognition during mating activity, acoustic traits have been widely used to clarify the taxonomy of anurans. They have been particularly useful in the study of taxa with high morphological similarity such as the Neotropical genus Physalaemus. Here, we reviewed the acoustic repertoires of the species of Physalaemus based on homology hypotheses in order to make comparisons more properly applicable for taxonomic purposes. We covered all the known clades and species groups for the genus, analyzing 45 species (94 % of the currently recognized taxa). Different call types were labeled with letters (i.e., A, B, and C) to avoid speculative functional propositions for the call types. In order to identify correctly the observed frequency bands, we propose a method to interpret them based on the predicted graphic behavior on audiospectrogram and on the mathematic relationship among bands considering each kind of band production (e.g., harmonics and sidebands). We found different acoustic traits between the major clades P. signifer and P. cuvieri. Species in the P. signifer clade have more than one call type (67 % of species in the clade). Furthermore, all species of this clade have A calls with pulses and/or low fundamental frequency (< 500 Hz). In the P. cuvieri clade, species emit only one call type and, in most species, this call is a continuous whine-like emission with relatively high fundamental frequency (> 400 Hz) and several S-shaped harmonics (except for species of P. henselii and P. olfersii groups, P. centralis, and P. cicada). Within the P. signifer clade, pulsed calls are present in P. angrensis, P. atlanticus, P. bokermanni, P. crombiei, P. irroratus, P. moreirae, P. nanus, and P. obtectus, whereas within the P. cuvieri clade this feature is restricted to a few species (10 % of the clade): P. jordanensis, P. feioi, and P. orophilus. A principal component analysis of the quantitative data indicates two clusters that substantially correspond to the composition of these two major clades with a few exceptions. Overall, the cluster composed of taxa of the P. signifer clade has lower fundamental frequency, bandwidth and dominant frequency at the end of the call and higher frequency delta and dominant frequency at the end of the call than the cluster with most taxa of the P. cuvieri clade. We also identified and described several similarities among acoustic signals of closely related species, which might correspond to synapomorphies in the evolution of the acoustic signal in the group. Species of the P. deimaticus group emit long sequences of very short A calls with low fundamental frequency (< 300 Hz) and short duration (< 0.2 s). Most species in the P. signifer group have clearly pulsed calls and emit at least two different call types. Species in the P. henselii group have calls with only high frequency bands (> 1700 Hz). Species in P. cuvieri group have continuous calls that resemble nasal-like sounds or whines, with downward frequency modulation. Species in the P. olfersii group emit long calls (> 1 s) with ascendant and periodic frequency modulation. Calls of the species in the P. biligonigerus and P. gracilis groups usually have continuous whine-like calls with call envelopes very variable within species. In addition, we describe traits in the genus for the first time, such as complex traits not predicted by simple and linear acoustic models (nonlinear phenomena), and discuss the application of acoustic traits to taxonomy and phylogenetics and morphological constraints of the vocal apparatus that might be related to the different acoustic properties found.

Physalaemus soaresi Izecksohn, 1965 We found a single call type for the species, referred to as call A. The call is composed of a single harmonic note with long duration, PFM, and a slight PAM, with no silence intervals. Bands have a general upward FM and a downward FM at the end, yielding arc-shaped bands in audiospectrogram when considering the entire call. Subharmonics, shifts of the fundamental frequency and deterministic chaos are common at the beginning and end of the calls. Call A (Fig. 38 A–J and 33F). We examined three recordings, a total of five minutes, with 40 calls from five males. Only some of these calls were measured (see Table 2). Call duration varies from 1.597 to 1.748 s. In most calls, the limits between the call rise, sustain, and fall are not clear. Usually, call rise and fall are similar in duration and shape (logarithmic). In some calls, the rise is longer than fall. There is a long sustain. It is usually regular, with a convex shape, but it can be almost flat (Fig. 38A, C), or have irregular AM segments, yielding amplitude peaks and valleys over the segment (Fig. 38D, E, F). The amplitude peak is usually at around the middle of the call duration. The envelope of the call varies between elliptic (Fig. 38A, C), rectangular (Fig. E, F), or triangular (pointed left; Fig. 38D), depending on the shape and steepness of the sustain. More than 50 % of the call energy is concentrated in 39 % of the call duration around the amplitude peak. Some calls have a slight PAM (there is no silence interval between peaks; Fig. 38F). The rate of the PAM is ca. 16 Hz, forming ca. 22 amplitude peaks throughout the call. The call has a harmonic series (Fig. 33F). The fundamental frequency is ca. 600 Hz. This band and the next harmonic are absent in the audiospectrogram. There are usually ca. six emphasized harmonics. Generally, the wave periods are regular and harmonics are clear throughout the call. However, subharmonics (f 0 1/2), jumps of the fundamental frequency, and deterministic chaos are common at the beginning and end of the call (Fig. 38B, G, H, I, J). The dominant frequency varies from ca. 2450 to 3060 Hz. The dominant harmonic varies from the second to the seventh, but it is usually the fourth or fifth (Fig. 33F). There is no clear shift in the relative energy between the bands throughout the call (Fig. 33F). Most of the call energy is between 2250 and 3750 Hz (three harmonics). The call has a general upward FM and a short downward FM at the end, yielding an arc-shaped bands in audiospectrogram when considering the entire call (Fig. 38B, G, H, I, J). Additionally, there is clear PFM throughout the call, which is usually independent from the PAM or can be directly proportional and synchronic to some parts of the PAM (Fig. 38 A–J)., Published as part of Hepp, Fábio & Pombal, José P., 2020, Review of bioacoustical traits in the genus Physalaemus Fitzinger, 1826 (Anura: Leptodactylidae: Leiuperinae), pp. 1-106 in Zootaxa 4725 (1) on page 70, DOI: 10.11646/zootaxa.4725.1.1, http://zenodo.org/record/3612996

Physalaemus cuvieri Fitzinger, 1826 We found a single call type for the species, referred to as call A. The call is composed of a single harmonic note, usually with a triangular envelope that resembles an arrow-like shape. It has a general downward FM, with an updownward FM segment in the first half of the call and a short upward FM segment at the end. Subharmonics are always present in the first half of the call. Call A (Fig. 32 A–V and 33A). We examined 75 recordings, a total of 94 minutes, with ca. 10200 calls from 228 males. Only some of these calls were measured (see Table 2). Call duration varies from 0.257 to 0.311 s. The envelope is very variable (Fig. 32A, C–G, M–Q). In most calls, the limits between the call rise, sustain and call fall are not clear. The ratio between call rise and fall duration, and their shape, are highly variable. Most calls have a fall longer than rise, or both have the same duration. Their shape varies from exponential to linear or logarithmic. The call rise has two consecutive exponential parts, the first shorter than the second. The sustain, when present, is irregular, usually composed of a shallow or deep valley (i.e., with a concave shape; Fig. 32A, C, D, G, M, Q). The amplitude peak is usually before the middle of the call duration. The envelope varies from elliptic (Fig. 32A, C, D, N, O, P, Q) to triangular (pointed right; Fig. 32E, M). Due to the concave shape of the sustain, the triangular envelope of some calls resembles an arrow. More than 50 % of the call energy is concentrated in 20 % of the call duration around the amplitude peak. There is no PAM in the call. The call has a harmonic series (Fig. 33A). The fundamental frequency is ca. 650 Hz and approximately the first seven harmonics are emphasized. The wave periods are regular and harmonics are clear throughout the call. Subharmonics (f 0 1/2) are present in ca. the first half of all calls examined (this part can be shorter or longer than one half of the call duration; Fig. 32B, H–L, R–V). The dominant frequency varies from ca. 690 to 780 Hz (Fig. 32B). The dominant harmonic is the first or second (at the very end of the call), but it is usually the first. There is a clear shift in the relative energy among bands. Although there is no shift in the dominant frequency, the higher bands get more energy toward the end of the call (Fig. 32K, L, T, V). Most of the call energy is between 500 and 1300 Hz (one or two harmonics). The call has a general downward FM (Fig. 32B, H–L, R–V). Additionally, the calls have an up-downward FM in the first half of the call duration, forming arc-shaped bands in this part of the call, and a short upward FM at the end (Fig. 32B, H–L, R–V). The general downward FM and the initial up-downward FM result in S-shaped harmonics when considering the entire call. There is no PFM., Published as part of Hepp, Fábio & Pombal, José P., 2020, Review of bioacoustical traits in the genus Physalaemus Fitzinger, 1826 (Anura: Leptodactylidae: Leiuperinae), pp. 1-106 in Zootaxa 4725 (1) on page 64, DOI: 10.11646/zootaxa.4725.1.1, http://zenodo.org/record/3612996

Physalaemus lisei Braun & Braun, 1977 We found a single call type for the species, referred to as call A. The call is composed of a single harmonic note with a long duration, slight PAM (no silence intervals between peaks) and irregular PFM. The bands have a general downward FM and a short upward FM segment at the end. Calls usually have nonlinear regimes such as deterministic chaos and subharmonics. Call A (Fig. 51 A–N and 52B). We examined 19 recordings, a total of 89 minutes, with ca. 915 calls from 50 males. Only some of these calls were measured (see Table 2). Call duration varies from 0.967 to 1.997 s. The envelope of the call is variable; durations of call rise and fall are usually short and similar in duration, with a long sustain in between; the rise and fall shapes vary from logarithmic to almost linear or exponential. The sustain is flat (Fig. 51E, F, H, I) or gradually ascending (Fig. 51C, G). Some calls have a final part with higher amplitude (Fig. 51C, G, I). Shallow and short amplitude valleys can be present, mainly at the beginning and end of the call (Fig. 51C, I). The amplitude peak is usually at the very end of the call duration. Depending on the slope of the sustain, the envelope varies from rectangular (Fig. 51E, F, H, I) to triangular (pointed left; Fig. 51C, G). More than 50 % of the call energy is concentrated in 47 % of the call duration around the amplitude peak. The call can have a slight PAM (there is no silence interval between peaks; Fig. 51E, G, H). The rate of the PAM is ca. 26 Hz, forming ca. 25 cycles throughout the call. The call has a harmonic series (Fig. 52B). The fundamental frequency is ca. 480 Hz and this band can be present with low energy or absent in audiospectrograms. Six adjacent harmonics are emphasized (first seven except the fundamental). Usually, the wave periods are regular and harmonics are clear throughout the call. However, several calls show nonlinear regimes such as subharmonics (f 0 1/2, f 0 1/3, f 0 1/4, or f 0 1/5), biphonation, and deterministic chaos (Fig. 51D, J, K, M). These phenomena can occur over the entire call. The dominant frequency varies from ca. 2330 to 2460 Hz (Fig. 51D). The dominant harmonic varies from the first to the fifth (except the second), but it is usually the fourth or fifth along the first half of the call (Fig. 52B). There is a clear shift in relative energy between the bands. Although there is no shift in the dominant frequency, higher bands get more energy towards the end of the call (Fig. 51D, L, N). Most of the call energy is between 950 and 3350 Hz (five to six harmonics). The call has a slight general downward FM (Fig. 51D, L, N). Additionally, calls have a very short and slight up-downward FM at their outset, leading to slightly arc-shaped bands in this part of the call, and a short upward FM at the end (Fig. 51L, N). The general downward FM and the initial up-downward FM result in S-shaped harmonics when considering the entire call. Some calls show clear PFM (Fig. 51L, N). Calls are usually emitted in irregular sequences, with two or three calls (Fig. 51 A–B)., Published as part of Hepp, Fábio & Pombal, José P., 2020, Review of bioacoustical traits in the genus Physalaemus Fitzinger, 1826 (Anura: Leptodactylidae: Leiuperinae), pp. 1-106 in Zootaxa 4725 (1) on page 82, DOI: 10.11646/zootaxa.4725.1.1, http://zenodo.org/record/3612996

Physalaemus cicada Bokermann, 1966 We found a single call type for the species, referred to as call A. The call is composed of a single harmonic note with an elliptic envelope and very short duration. It has a general downward FM, with an up-downward FM segment in the first half of the call (Fig. 36B, F). Calls are emitted in long sequences (more than 300 calls per sequence; Fig. 36C, D). Call A (Fig. 36 A–G and 33D). We examined six recordings, a total of 13 minutes, with ca. 12500 calls from eight males. Only some of these calls were measured (see Table 2). Call duration varies from 0.004 to 0.047 s. The call rise and fall are similar in duration and shape (exponential). The sustain is short or absent. The envelope is elliptic (Fig. 36A. E). The amplitude peak is at around the end of the first two fifths of the call duration. More than 50 % of the call energy is concentrated in 24 % of the call duration around the amplitude peak. There is no PAM in the call. The call has a harmonic series (Fig. 33D). The fundamental frequency is ca. 410 Hz and approximately the first ten harmonics (except the first one) are emphasized. The wave periods are regular and harmonics are clear throughout the call. The dominant frequency varies from ca. 1410 to 3560 Hz (Fig. 33D, 36B). The dominant harmonic varies from the seventh to the 10 th, but it is usually the eighth. There is a clear shift in relative energy between bands; the dominant frequency gets higher toward the end of the call, starting at the seventh or eighth harmonic and ending at the eighth, ninth or 10 th (Fig. 33D, 36F). Most of the call energy is between 950 and 3850 Hz (eight harmonics). The call has a general downward FM (Fig. 36B, F). Additionally, calls have an up-downward FM in the first half of the call duration, yielding arc-shaped bands in this part of the call, and a short downward FM at the end (Fig. 36B, F). The general downward FM and the initial up-downward FM result in S-shaped harmonics when considering the entire call (Fig. 36F). There is no PFM. Calls are usually emitted in series, resulting in long call sequences of ca. 400 calls in each sequence (Fig. 36C, D, E, F)., Published as part of Hepp, Fábio & Pombal, José P., 2020, Review of bioacoustical traits in the genus Physalaemus Fitzinger, 1826 (Anura: Leptodactylidae: Leiuperinae), pp. 1-106 in Zootaxa 4725 (1) on page 68, DOI: 10.11646/zootaxa.4725.1.1, http://zenodo.org/record/3612996

Physalaemus lateristriga (Steindachner, 1864) We found a single call type for the species, referred to as call A. The call is composed of a single harmonic note with a long duration and a slight PAM (no silence intervals between pulses). It has irregular and strong PFM throughout the call. The bands have no general FM or only slight general FM, which is usually upward. Call A (Fig. 43 A–F and 42B). We examined seven recordings, a total of 20 minutes, with ca. 160 calls from 16 males. Only some of these calls were measured (see Table 2). Call duration varies from 1.330 to 3.746 s. The call rise is restricted to the very beginning of the call, most of the call corresponding to the sustain (Fig. 43C). Call rise and fall are very short and similar to each other in duration. Sustain usually regular and almost flat (Fig. 43C), but some calls have convex or inclined segments, with amplitude gradually increasing towards its end (Fig. 43A, D). In some calls, there is a shallow valley at the beginning of the sustain (Fig. 43A, C). The amplitude peak is at around the middle or at the end of the call duration. The envelope of the call varies from rectangular (Fig. 43C) to triangular (pointed left; Fig. 43A, D). More than 50 % of the call energy is concentrated in 39 % of the call duration around the amplitude peak. The call has a slight PAM (silence intervals absent between peaks; Fig. 43A, D). The rate of the PAM is ca. 8 Hz, forming ca. 29 cycles throughout the call. The cycle rise and fall are similar and the amplitude peak is at the middle of the cycle duration. The call has a harmonic series (Fig. 42B). The fundamental frequency is ca. 170 Hz. The first five harmonics are usually absent in the audiospectrogram or with very low energy. There are ca. eight adjacent emphasized harmonics. The wave periods are regular and harmonics are clear throughout the call. Subharmonics (f 0 1/2) are present at the beginning of some calls (Fig. 43B, F). The dominant frequency varies from ca. 1590 to 1840 Hz (Fig. 43B). The dominant harmonic varies from the ninth to the 13 th, but it is usually the ninth or 10 th (Fig. 42B). There is no clear shift in the relative energy between bands throughout the call. Most of the call energy is between 1100 and 2150 Hz (seven harmonics). Calls usually lack a clear general FM (Fig. 43B, E). In some calls, a slight up or downward general FM is observed, usually upward. A short downward FM is frequently present at the end of the call (Fig. 43E). Additionally, there is a strong PFM throughout the call, which is usually independent (Fig. 43C, E), but it is directly proportional and synchronic to PAM when it is present (Fig. 43A, B)., Published as part of Hepp, Fábio & Pombal, José P., 2020, Review of bioacoustical traits in the genus Physalaemus Fitzinger, 1826 (Anura: Leptodactylidae: Leiuperinae), pp. 1-106 in Zootaxa 4725 (1) on page 73, DOI: 10.11646/zootaxa.4725.1.1, http://zenodo.org/record/3612996

Physalaemus kroyeri (Reinhardt & Lütken, 1862) We found a single call type for the species, referred to as call A. The call is composed of a single harmonic note with a general downward FM throughout the call and an up-downward FM segment in the first fifth of the call duration. Call A (Fig. 30 A–F and 24G). We examined four recordings, a total of six minutes, with ca. 190 calls from eight males. Only some of these calls were measured (see Table 2). Call duration varies from 0.673 to 0.759 s. In some calls the limits between the call rise, sustain and call fall is not very clear (mainly in calls with elliptic envelope, Fig. 30A). The call rise and fall are similar in duration. Usually, the call rise has a short logarithmic-shaped section followed by an exponential shape, whereas call fall has an exponential shape only. The sustain is irregular, usually with short or long shallow valleys (Fig. 30D). The amplitude peak is usually before the middle of the call duration. The envelope varies from elliptic (Fig. 30A, C) to almost rectangular (when flat sustains are present; Fig. 30D). More than 50 % of the call energy is concentrated in 34 % of the call duration around the amplitude peak. There is no PAM in the call. The call has a harmonic series (Fig. 24G). The fundamental frequency is ca. 480 Hz and approximately the first seven harmonics are emphasized. The wave periods are regular and harmonics are clear throughout the call. The dominant frequency varies from ca. 2060 to 2160 Hz (Fig. 30B). The dominant harmonic varies from the second to the sixth (Fig. 24G, 30E, F). There is a clear shift in the relative energy between the bands; the dominant frequency gets higher until three fourths of the call duration, starting at the second harmonic and ending at the sixth; thenceforth, it gets lower, ending at the third harmonic (Fig. 24G, 30E, F). Most of the call energy is between 450 and 2700 Hz (four to seven harmonics). The call has a general downward FM (Fig. 30B, E, F). Additionally, the calls have an up-downward FM in the first fifth of the call duration, yielding arc-shaped bands in this part of the call, and a short downward FM at the end (Fig. 30B, E, F). The general downward FM and the initial up-downward FM result in S-shaped harmonics when considering the entire call. There is no PFM., Published as part of Hepp, Fábio & Pombal, José P., 2020, Review of bioacoustical traits in the genus Physalaemus Fitzinger, 1826 (Anura: Leptodactylidae: Leiuperinae), pp. 1-106 in Zootaxa 4725 (1) on page 62, DOI: 10.11646/zootaxa.4725.1.1, http://zenodo.org/record/3612996