A New Dwarf Species of Proceratophrys Miranda-Ribeiro, 1920 (Anura, Cycloramphidae) from the Highlands of Chapada Diamantina, Bahia, Brazil

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A New Dwarf Species of Proceratophrys Miranda-Ribeiro, 1920 (Anura, Cycloramphidae) from the Highlands of Chapada Diamantina, Bahia, Brazil Zootaxa 3551: 25–42 (2012) ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ ZOOTAXA Copyright © 2012 · Magnolia Press Article ISSN 1175-5334 (online edition) urn:lsid:zoobank.org:pub:BC6CB247-BCCA-49FD-A014-E3A5FDB366DC A new dwarf species of Proceratophrys Miranda-Ribeiro, 1920 (Anura, Cycloramphidae) from the highlands of Chapada Diamantina, Bahia, Brazil MAURO TEIXEIRA JUNIOR1,2, RENATA CECÍLIA AMARO1, RENATO SOUSA RECODER1, FRANCISCO DAL VECHIO1 & MIGUEL TREFAUT RODRIGUES1 1Laboratório de Herpetologia, Departamento de Zoologia, Instituto de Biociências, Rua do Matão, no 101, CEP 05508-090, Universi- dade de São Paulo, São Paulo, Brazil 2Corresponding author. E-mail: [email protected] Abstract A new species of Proceratophrys is described from the highlands of northeastern Brazil. Molecular and morphological data suggests that Proceratophrys redacta sp. nov. is sister to P. minuta, and related to P. schirchi and P. cristiceps. The new species is diagnosed by its small size, absence of rostral and palpebral appendages, sagittal ridges interrupted, absence of postocular swellings, snout vertical in profile and dorsal coloration lacking distinct ocelli. The new species represents another example of endemism for the genus Proceratophrys in Chapada Diamantina region, and of another appendageless small-sized species associated with highlands. The phylogenetic results indicate that current morphological groupings in Proceratophrys may not represent natural groups. Key words: Proceratophrys redacta sp. nov., mountain endemism, frog diversity Introduction After his travel through the Brazilian coast, the German naturalist Alexander Phillip Maximilian, Prinz zu Wied- Neuwied (see Myers et al. 2011), described the horned frog, Ceratophrys boiei (Wied 1824), which underwent taxonomic rearrangement (Gravenhorst (1825; 1829), ending up to be the first described species in the currently recognized genus Proceratophrys. After that several species were described (Peters 1872; Günther 1873; Müller 1883; Miranda-Ribeiro 1920; 1926; 1937; Jim & Caramaschi 1980; Izecksohn & Peixoto 1981; Weygoldt & Peixoto 1985; Giaretta & Sazima 1993; Mercadal de Barrio & Barrio 1993; Eterovick & Sazima, 1998; Izecsohn et al. 1999; Giaretta et al. 2000; Kwet & Faivovich 2001; Cruz et al. 2005; Prado & Pombal 2008; Cruz & Napoli 2010; Frost 2011; Martins & Giaretta 2011; Napoli et al. 2011; Cruz et al. 2012) and its taxonomy rearranged (Lynch 1971; Prado & Pombal 2008; Amaro et al. 2009). Currently the genus comprises 27 species of small to medium/large-sized frogs, occurring in forests, savannas, grasslands and shrubby lands of eastern, central and southern South America, in Brazil, Argentina and Paraguay (Prado & Pombal 2008; Frost 2011; Martins & Giaretta 2011; Napoli et al. 2011). However, its present richness is likely underestimated, as indicated by the constant discovery of new species (Prado & Pombal 2008; Ávila et al. 2011; Martins & Giaretta 2011; Napoli et al. 2011; Cruz et al. 2012). Furthermore, preliminary molecular phylogenies are showing that species formerly attributed to Odontophrynus, are indeed members of Proceratophrys (Amaro et al. 2009). Recently, a dwarf species of Proceratophrys (P. m inuta Napoli, Cruz, Abreu & Del Grande 2011) was described from the highlands of Chapada Diamantina, at the northern portion of Espinhaço range in state of Bahia (Napoli et al. 2011). This small-sized highland species seems to be restricted to the cloud forest patches of high elevations (above 800 m a.s.l., being very common at about 1170 m a.s.l.). Based on similarities in external morphology this new species was supposed to be allied to P. schirchi, (Napoli et al. 2011). Both P. s c h irc hi and P. m in ut a are not currently assigned to any species group, as well as P. rondonae, however the remaining species have been informally assigned to three or four species groups, based on overall Accepted by M. Vences: 10 Oct. 2012; published: 16 Nov. 2012 25 morphological similarity. The P. boiei species complex, characterized by a single and long uni-cuspidate palpebral appendage and lack of a triangular rostral appendage, is thought to include P. b oi e i (Wied, 1824); P. pa v io ti i Cruz, Prado and Izecksohn, 2005; and P. re n al is (Miranda-Ribeiro, 1920). The P. appendiculata species complex, sometimes referred to as a subset of P. bo ie i group, also with a single and long uni-cuspidate palpebral appendage, additionally characterized by the presence of a triangular rostral appendage: P. appendiculata (Günther, 1873); P. laticeps Izecksohn and Peixoto, 1981; P. melanopogon (Miranda- Ribeiro, 1926), P. m o e h r in gi Weygoldt and Peixoto, 1985; P. phyllostomus Izecksohn, Cruz and Peixoto, 1998; P. sanctaritae Cruz and Napoli, 2010; P. subguttata Izecksohn, Cruz and Peixoto,1998; and P. tupinamba Prado and Pombal, 2008. Four species have been associated to the P. bigibbosa group, characterized by the presence of postocular swellings (Kwet & Faivovich 2001): P. avelinoi Mercadal de Barrio & Barrio, 1993; P. bigibbosa (Peters, 1872); P. brauni Kwet & Faivovich, 2001; and P. palustris Giaretta & Sazima, 1993. Finally, eight species have been associated with the P. cristiceps group, which gathers species lacking both prominent supraocular appendages and postocular swellings, but does not have any recognized shared derived characters (Giaretta et al. 2000): P. ar i du s Cruz, Nunes and Junca, 2012; P. caramaschi Cruz, Nunes and Junca, 2012: P. cristiceps (Müller, 1883); P. concavitympanum Giaretta, Bernarde & Kokubum, 2000; P. cururu Eterovick & Sazima, 1998; P. goyana (Miranda-Ribeiro, 1937); P. moratoi (Jim and Caramaschi, 1980); P. strussmannae Ávila, Kawashita-Ribeiro & Morais, 2011; and P. vielliardi Martins & Giaretta, 2011. Herein, based on a series of specimens recently collected at the highlands of northern Chapada Diamantina, at Morro do Chapéu municipality, we describe a new dwarf species of Proceratophrys providing data on its natural history, its phylogenetic placement, as well as those for P. minuta, discussing the evolution of body shape and the validity of the recognized species groups within Proceratophrys. Material and methods Field sampling. Fieldwork was carried out during the rainy season, from 22nd December 2011 to 2nd January 2012, at Morro do Chapéu municipality. Ten pitfall traps with drift fences were installed at a plain sandy Caatinga, and 25 others at a montane forest at the slopes of Morro do Chapéu hill. Unquantified active search was also performed at the surrounding areas and at Ferro Doido waterfall. Collected specimens were killed with lethal doses of anesthesics, fixed in 10% formalin, preserved in 70% alcohol and housed at the herpetological collection of Museu de Zoologia da Universidade de São Paulo, São Paulo, Brazil. Morphological assessment. Morphological comparisons were based on voucher specimens housed at the herpetological collection of the Museu de Zoologia da Universidade de São Paulo (MZUSP) and specimens collected by the Laboratório de Herpetologia, Instituto de Biociências da Universidade de São Paulo (MTR) (Appendix I), and data available in the literature (Giaretta & Sazima 1993; Mercadal de Barrio & Barrio 1993; Eterovick & Sazima 1998; Giaretta et al. 2000; Kwet & Faivovich 2001; Cruz et al. 2005; Prado & Pombal 2008; Cruz & Napoli 2010; Santana et al. 2010; Ávila et al. 2011; Martins & Giaretta 2011; Napoli et al. 2011; Cruz et al. 2012). Diagnostic features and morphological descriptions follow Prado and Pombal (2008) and Napoli et al. (2011). Measurements were taken using a digital caliper to the nearest 0.01 mm and rounded to the first decimal unit to avoid pseudoprecision (Hayek et al. 2001). Twelve morphometric measurements were taken following Heyer et al. (1990) and Prado and Pombal (2008): snout-vent length (SVL); head length (HL); head width (HW); eye diameter (ED); eye to nostril distance (END); upper eyelid width (UEW); interorbital distance (IOD); internostril distance (IND); thigh length (THL); tibia length (TL); foot plus tarsus length (FTL); and hand length (HAL). All individuals were measured, as all were consider adults: males with vocal slits and females with vitelogenic eggs visible though skin. Moreover individuals were found in amplexus, attesting for their maturity. Geographical distribution. The geographical distribution of Proceratophrys was assessed, to present the distribution of the new species in the genus context and to add some data to the discussion on validity of some species. For this purpose records were compiled, mostly from literature (Miranda-Ribeiro 1955; Jim & Caramaschi 1980; Giaretta & Sazima 1993; Mercadal de Barrio & Barrio 1993; Rossa-Feres & Jim 1996; Caramaschi & Velosa 1997; Eterovick & Sazima 1998; Machado et al. 1999; Giaretta et al. 2000; Brandão & Araujo 2001; Giasson et al. 2001; Kwet & Faivovich 2001; Boquimpani-Freitas et al. 2002; de Sá & Langone 2002; Lavilla et al. 2002; Kwet & Baldo 2003; Silvano & Pimenta 2003; Hiert & Moura 2004; Pimenta & Carvalho-e-Silva 2004; Cruz et al. 2005; 26 · Zootaxa 3551 © 2012 Magnolia Press TEIXEIRA JR. ET AL. Brusquetti & Lavilla 2006; Leite et al. 2006; Cacciali et al. 2007; Vieira et al. 2007; de Lima 2007; Zanella & Busin 2007; Brasileiro et al. 2008; Gonsales 2008; Lucas & Foster 2008; Prado & Pombal 2008; Vieira et al. 2008; Amaro et al. 2009; Cintra et al. 2009; Oda et al. 2009; Santos et al. 2009; Caldart et al. 2010; Carosini et al. 2010; Carvalho et al. 2010; Cruz & Napoli 2010; Fatorelli et
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