A new species of Pseudopaludicola Miranda-Ribeiro (Leiuperinae: Leptodactylidae: Anura) from the Cerrado of southeastern Brazil Author Andrade, Felipe Silva De Author Carvalho, Thiago Ribeiro De text Zootaxa 2013 3608 5 389 397 journal article 10.11646/zootaxa.3608.5.7 97a20516-6489-4f57-92f1-40bf8dee7951 1175-5326 219280 51879195-F141-4FFF-9AA9-9394FD650EC7 Pseudopaludicola facureae , new species ( Figures 2–3 ) Holotype : AAG-UFU 0 853, adult male, collected at the Clube Caça e Pesca Itororó de Uberlândia ( 18°58'30.46"S , 48°17'26.23"W ; 790 m above sea level), Municipality of Uberlândia, State of Minas Gerais, southeastern Brazil , in October 2011 by A.A. Giaretta. Paratypes : Two adult male specimens collected at the Clube Caça e Pesca Itoror Uberlândia, Municipality of Uberlândia, State of Minas Gerais, Brazil : AAG-UFU 0854–0855, collected by A.A. Giaretta in October 2011 . Twenty adult specimens in the Jardim Karaíba neighborhood, Municipality of Uberlândia, State of Minas Gerais, Brazil : ten males: ZUEC 13650 collected by A.P. Rodrigues and A.A. Giaretta in March 2006 ; ZUEC 13651 collected by A.A. Giaretta on 24 February 2001 ; ZUEC 13652 collected by A.A. Giaretta on 25 November 2000 ; AAG-UFU 2281 collected by A.A. Giaretta on 28 February 2001 ; AAG-UFU 2277–2278 collected by A.A. Giaretta on 0 9 February 2001 ; AAG-UFU 2528 collected by A.A. Giaretta and K.G. Facure on 0 9 October 2003 ; AAG-UFU 2622 collected by A.A. Giaretta on 0 6 March 2004 ; AAG-UFU 3586 collected by A.A. Giaretta on 25 November 2000 ; AAG-UFU 3588 collected by A.A. Giaretta and K.G. Facure on 29 October 2000 . Females: ZUEC 13653 collected by A.A. Giaretta on 27 September 2001 ; ZUEC 13654, AAG-UFU 3587 collected by A.A. Giaretta on 24 February 2001 ; AAG-UFU 1160 collected by A.A. Giaretta and F.S. Andrade on 0 1 June 2012 ; AAG-UFU 2279 collected by A.A. Giaretta on 0 9 February 2001 ; AAG-UFU 2282 collected by A.A. Giaretta on 28 February 2001 ; AAG-UFU 3585 collected by A.A. Giaretta on 25 November 2000 ; AAG-UFU 3589 collected by A.A. Giaretta in June 2000 ; AAG-UFU 3591 collected by A.P. Rodrigues and D.R. Silva on 0 7 May 2006 ; AAG-UFU 4731 collected by A.A. Giaretta, M.N.C. Kokubum, M. Menin, and R. Alvarenga on 31 October 2000 . Juvenile specimens: AAG-UFU 1157–1159 , and AAG-UFU 1161–1164 collected by A.A. Giaretta and F.S. Andrade on 0 1 June 2012 . Referred specimens : Pseudopaludicola aff. canga — Brazil , Minas Gerais, Uberlândia: AAG-UFU 2608–2609 (Giaretta & Facure 2009); ZUEC 14181, 14185–14187, 14209, 14212, 14214, 14216–14217, 14222–14223 (Duarte et al. 2010). Pseudopaludicola sp.— Brazil , Minas Gerais, Uberlândia: AAG-UFU 4728–4732 (Carvalho 2012). Diagnosis. Pseudopaludicola facureae sp. nov. is assigned to the genus by possessing hypertrophied antebrachial tubercle. The new taxon is diagnosed by the following combination of characters: (1) small size (SVL 12.1–15.1 mm in adult males); (2) absence of either T-shaped terminal phalanges or expanded toe tips (disks or pads); (3) absence of enlarged palpebral tubercles; (4) short hindlimbs (tibiotarsal articulation reaching the eye); (5) advertisement call composed of series of non-pulsed notes, emitted in well-defined sequences. Comparisons with other species. Pseudopaludicola facureae sp. nov is promptly diagnosed from the P. pusilla species group by the absence of either T-shaped terminal phalanges or expanded toe tips (disks or pads). The new species can also be distinguished from P. boliviana , P. ceratophyes , and P. llanera by the absence of enlarged palpebral tubercle (Lynch 1989). Pseudopaludicola facureae sp. nov. ( 12.1–15.1 mm in adult males) is diagnosed from P. giarettai , P. riopiedadensis , P. saltica , P. serrana , and P. ternetzi (combined adult male SVL 15.0– 19.7 mm ) (see Table 2 in Carvalho 2012) by its smaller snout-vent length. Morphological/morphometric comparisons with P. riopiedadensis were very limited due to the lack of data on the species in the original description (see Mercadal de Barrio & Barrio 1994). Pseudopaludicola facureae sp. nov. is distinguished from P. m u r u n d u , P. saltica , and P. serrana by having short hindlimbs (tibiotarsal articulation reaching the eye), whereas all three abovementioned species have long hindlimbs (tibiotarsal articulation extending beyond the tip of snout). Pseudopaludicola facureae sp. nov. is diagnosed from almost all congeners by possessing the advertisement call (fig. 4) composed of series of non-pulsed notes in comparison with that of congeners [pulsed advertisement calls: P. boliviana (Duré et al. 2004) , P. saltica and P. falcipes (Haddad & Cardoso 1987) , P. mystacalis (A. Pansonato unpubl. data), P. murundu (Toledo et al. 2010) , P. serrana (Toledo 2010) , P. mineira (Pereira & Nascimento 2004) , P. riopiedadensis (L.D. Vizotto pers. comm.; A. Pansonato unpubl. data), and P. ternetzi (A. Pansonato unpubl. data)]. From P. canga , P. giarettai , and P. hyleaustralis , all three possessing non-pulsed advertisement calls, the new species is diagnosed by the emission of the advertisement call in note series, note duration, note rate per minute, and dominant frequency. The advertisement call of P. fa c ure a e sp. nov. (fig. 4) is composed of 3–53 notes/call, whereas topotypic P. canga releases series of up to nine notes (Giaretta & Kokubum 2003); and non-topotypic populations releases series of up to 19 notes (Pansonato et al. 2012). P. giarettai releases isolated long (117–187 ms) notes (Carvalho 2012), whereas the new species releases short (15–35 ms) notes in series. Note duration of P. hyleaustralis ranges from 25–50 ms, emitted at a rate of 504–623 notes/minute, and dominant frequency from 3605–4164 Hz (Pansonato et al. 2012), whereas P. f a c u re a e sp. nov. has a shorter note duration (15–35 ms), a higher emission rate (480–1860 notes/minute), and a higher dominant frequency (4076–5108 Hz). Pseudopaludicola facureae sp. nov. (2n = 18 chromosomes; see P. aff. canga from Uberlândia in Duarte et al. 2010) is also diagnosed from P. m ys t ac a li s (2n = 16), P. falcipes , P. m i n e i r a , P. murundu , and P. saltica (2n = 22), and P. ternetzi (2n = 20), by a distinctive chromosome number (Duarte et al. 2010; Toledo 2010; Fávero et al. 2011). Description of holotype . Snout subovoid from above, rounded in lateral view ( sensu Heyer et al. 1990) (figs. 3A-B). Nostrils closer to the snout tip than to the eyes; pupil rounded; upper eyelids. Canthus rostralis rounded, smooth; single subgular vocal sac; choanae well-separated from each other; vocal slits present. Vomerine teeth absent; tongue ovoid, free posteriorly; no pigmentation on the base of tongue. Tympanic ring undefined; lateral of head and flanks with discrete granules (superior surface of limbs). One ovoid antebrachial tubercle present in the first quarter of the forearm; outer metacarpal tubercle round; inner metacarpal tubercle elongated; subarticular tubercles rounded, supernumerary tubercles indistinct; fingers extensively fringed; outer ridge from the outer metacarpal tubercle to almost the tip of finger IV (fig. 3C); relative length of fingers I<II~IV<III; inner metatarsal tubercle oval; outer metatarsal tubercle round, 1/3 of the size of the inner metatarsal tubercle; outer metatarsal tubercle conical (fig. 3D); relative length of toes I<II<III~V<IV; finger and toe tips not expanded laterally; a dermal ridge from the inner metatarsal tubercle to 1/2 of the length of tarsus ending in a discrete, enlarged tubercle; outer fringe of toe V extending to almost the outer metatarsal tubercle; discrete fine nuptial asperity present on the base of thumbs; toes webbed basally and extensively fringed to almost their tips; outer edge of tarsus and forearm, and heel, smooth. Cloacal region smooth. Belly and ventral limb surfaces smooth. A well-defined vertebral stripe from the snout tip to vent; transverse stripes on thighs (2–3), shanks (3–4), feet (3–4), and forearms (2–3). Dorsal surfaces smooth, with some scattered enlarged granules on shanks; a broad discontinuous stripe on the posterior surface of thighs, passing below vent. Measurements of holotype (mm). SVL 13.6, HL 3.5, HW 4.6, IND 1.1, END 1.2, ED 1.6, HAL 4.0, TL 7.2, SL 7.3, FL 7.8. Color of holotype in preservative. In preservative (fig. 2), grayish brown surfaces in dorsal view, with darker brown blotches and a white vertebral stripe. Belly unpigmented (whitish); ventral surface of hands and feet dark gray, mottled. Vocal sac with a few dark brown mottled; posterior surface of thighs with brown blotches and with a cream stripe. FIGURE 1. Paratopotypes of Pseudopaludicola facureae sp. nov. in life. A—adult female (AAG-UFU 1157; SVL = 16.1 mm); B—juvenile female (AAG-UFU 1158; SVL = 13.4 mm); C—juvenile male (AAG-UFU 1164; SVL 14.8 mm); D—adult female (AAG-UFU 1160; SVL = 15.3 mm). Specimens from the Municipality of Uberlândia, State of Minas Gerais, Brazil. FIGURE 2. Pseudopaludicola facureae sp. nov. , adult male, holotype (AAG-UFU 0853). (A) Dorsal and (B) ventral views. SVL = 13.6 mm. TABLE 1. Morphometric characters (in millimeters) of the Pseudopaludicola facureae sp. nov. type series adult specimens (including the holotype) from the Municipality of Uberlândia, State of Minas Gerais, Brazil. Mean+SD (minimum–maximum). N = number of specimens analyzed.

Characters	Males N = 13	Females N = 10
SVL	13.2±0.9 (12.1–15.1)	16.1±1.1 (14.7–18.3)
HL	3.3±0.2 (3.0–3.8)	4.4±0.5 (3.9–5.3)
HW	4.7±0.3 (4.2–5.3)	5.6±0.4 (4.8–6.2)
ED	1.5±0.1 (0.9–1.4)	1.8±0.2 (1.6–2.0)
END	1.1±0.1 (0.9–1.4)	1.3±0.2 (1.2–1.8)
IND	1.1±0.1 (0.9–1.3)	1.4±0.2 (1.1–1.7)
HAL	3.9±0.3 (3.4–4.5)	4.5±0.3 (4.2–5.3)
TL	6.6±0.6 (5.6–7.6)	7.5±0.3 (7.0–7.8)
SL FL	7.0±0.6 (6.1–8.0) 7.7±0.5 (7.0–9.0)	8.3±0.3 (7.8–8.7) 9.2±0.4 (8.7–9.8)

Color in life . Dorsum dark gray or brown; flanks pale gray or brown; belly cream; pale cream blotches on a whitish background on chin; pale cream blotches on the lateral of head; a cream stripe on the posterior surface of thighs. Some specimens have a mottled vocal sac. Variation. The specimens AAG-UFU 0854–0855, 2528, 2277, 3586, and ZUEC 13650, 13652 have a discrete tympanic ridge from behind the eyes to the proximal portion of the arms. The specimens AAG-UFU, 3585, 3589, ZUEC 13650–13652, 13654 have paravertebral chevron-shaped dermal ridges from behind the eyes to the scapular region. The specimens AAG-UFU 1159, 2277–2279 , 2281–2282 , 2528, 2622, 3585–3588, 4731, and ZUEC 13650–13652 have dorsum and dorsal surfaces of limbs with some degree of discoloration in preservative. The specimens AAG-UFU 2277–2278 , 2281, 2622, 3586, 3588, and ZUEC 13650–13652 have the ventral surface of hands and feet dark brown, mottled. The specimens AAG-UFU 1158, 1164, 0854–0855, 2281–2282 , 2528, 3586–3588, 3591, and ZUEC 13652–13653 have no vertebral stripe. The specimen AAG-UFU 1164 had a green vertebral stripe in life. The specimens AAG-UFU 2528, and ZUEC 13651–13652 have the vocal sac with many warts. The females have a more robust body, nuptial pads absent. Natural history. A detailed description of P. f a c u re a e sp. nov. natural history aspects, including call activity, reproductive ecology, and tadpole description is found in Giaretta and Facure 2009 (referred to as Pseudopaludicola aff. canga ). Distribution. Pseudopaludicola facureae sp. nov. is known from the type locality (Municipality of Uberlândia), as well as in the Municipality of Limeira do Oeste (vocalizations; L.B. Martins), State of Minas Gerais, approximately 250 km in a straight line westward from the type locality. Etymology. The name is a noun in the genitive case honoring Kátia G. Facure, the first researcher who called attention to the occurrence of this species in the study region in October 2000 . FIGURE 3. Pseudopaludicola facureae sp. nov. , adult male, holotype (AAG-UFU 0853). (A) Dorsal and (B) lateral views of head; (C) ventral views of hand and (D) foot. Scale bar = 5 mm. Advertisement call. Ten males recorded (N = 88 advertisement calls). Quantitative variables are summarized in Table 2 . Advertisement call (fig. 4) consists of series of 3–53 non-pulsed notes/call (mean 16.0; SD = 3.5), emitted in sequences of 3–26 advertisement calls/sequence (mean 9.2; SD = 4.8); sequence duration was 3–22 seconds (mean 9.8; SD = 6.3). Advertisement call sequence was emitted at a rate of 1–2 sequences/minute (mean 1.2; SD = 0.4). Advertisement call duration was 21–3013 ms (mean 658.5; SD = 194.2), and intercall duration was 204–931 ms (mean 462.2; SD = 108.8). Advertisement call was emitted at a rate of 1–2 calls/second (mean 1.1; SD = 0.2). Note duration was 15–35 ms (mean 23.1; SD = 3.7), and internote duration was 12–104 ms (mean 27.0; SD = 9.8), with a rate of 8–31 notes/second (mean 17.0; SD = 2.5), and a rate of 480–1860 notes/minute (mean 1020.6; SD = 147.6). Dominant frequency was 4076–5108 Hz (mean 4500; SD = 263). TABLE 2. Bioacoustic variables of Pseudopaludicola facureae sp. nov. from the Municipality of Uberlândia, State of Minas Gerais, Brazil. Mean+SD (minimum-maximum). N = number of specimens recorded (number of analyzed advertisement calls).

Var i abl es Advertisement call duration (ms)	Pseudopaludicola facureae sp. N =10 (88) 658.5±194.2 (21–3013)	nov.
Intercall interval (ms)	462.2±108.8 (204–931)
Notes/advertisement call	16.0±3.5 (3–53)
Note duration (ms)	23.1±3.7 (15–35)
Internote duration (ms) Advertisement call sequence (s) Advertisement calls/sequence	27.0±9.8 (12–104) 9.8±6.3 (3–22) 9.2±4.8 (3–26)
Advertisement calls sequence/minute	1.2±0.4 (1–2)
Advertisement calls/second	1.1±0.2 (1–2)
Notes/second	17.0±2.5 (8–31)
Notes/minute	1020.6±147.6 (480–1860)
Dominant frequency (Hz)	4500±263 (4076–5108)

Before the emission of the advertisement call sequences, isolated notes with irregular structure, duration, and interval are emitted, herein referred to as introductory notes (fig. 4). Duration of introductory note sequence was 0.5–2.5 seconds (mean 1.7; SD = 0.5), with 4–15 notes/sequence (mean 10.4; SD = 2.7). Interval between introductory note sequences and advertisement call sequences was 74–317 ms (mean 165.1; SD = 75.1). FIGURE 4. A—Oscillogram (14 seconds) of 15 introductory notes followed by an entire advertisement call sequence (6 advertisement calls) of Pseudopaludicola facureae sp. nov. from the Municipality of Uberlândia, Minas Gerais, Brazil. B—Audiospectrogram (above) and corresponding oscillogram (below) detailing 9 introductory notes outlined by a blue rectangle. C—Audiospectrogram (above) and corresponding oscillogram (below) detailing the last advertisement call (26 notes) of the entire sequence outlined by a red rectangle. Sound file: Pseudop_facureaeUberlândiaMG9aAAGm671; approximately 17:30h, 28 October 2011; air 27ºC, water 25ºC. Vouchered recording. Remarks. Three populations from southeastern and northeastern Brazil were assigned to Pseudopaludicola aff. canga (Giaretta & Facure 2009; Duarte et al. 2010) on the basis of common chromosome number (2n = 18) and putative morphological relatedness. The population from Uberlândia, State of Minas Gerais, actually represents P. facureae , easily diagnosed from P. canga by a bioacoustic approach. These species are separated by a vertical distance of about 1500 km from each other considering their respective type localities. The population assigned to P. aff. canga from Icém (State of São Paulo), southeastern Brazil , occurs about 185 km southward from P. f a c u re a e type locality, and currently represents an additional undescribed species with distinctive morphological and bioacoustic evidence in comparison with both P. canga and P. f ac u re ae (Pansonato pers. comm.). The third population assigned to P. aff. canga from Barreirinhas (State of Maranhão), northeastern Brazil , was reassessed in Pansonato et al. (2012) and currently represents a new distributional record of P. c a n g a instead. Acknowledgements Grants by Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) (TRC) and Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) (FSA). Financial support to our laboratory came from Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) and Fundação de Amparo à Pesquisa do Estado de Minas Gerais (FAPEMIG). Special thanks go to Ariovaldo A. Giaretta, coordinator of our lab, and Lucas B. Martins for providing bioacoustic data on the newly described species; André Pansonato and Luiz Dino Vizotto for making available essential data on Pseudopaludicola , Vanessa Suzuki Kataguiri for providing laboratory facilities, and Isabelle Aquemi Haga for helping at laboratory.