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Bokermannohyla sapiranga sp. nov. (Figs. 1���2) Holotype. Brazil, Distrito Federal, Reserva Ecol��gica do Roncador (15 �� 55 ��� 49 ��� S, 47 �� 52 ��� 59 ��� W, 1110m asl), CHUNB 62384, adult male, R.A. Brand��o, R.D. Fran��oso, T. Marques, R.M. Japiassu, and L.R.A. Braga col., 10 December 2009. Paratypes. Brazil, Distrito Federal, Bras��lia (15 �� 52 ��� 25 ��� S, 47 �� 51 ��� 59 ��� W, 1070m asl), CFBH 2248���2249, adult males, A. Sebben col., no date. CHUNB 14383���14384, adult females, A. Sebben col., 0 1 April 1986. CHUNB 38832, adult male, unknown collector, 0 1 May 1993. CHUNB 14377���14379, juveniles, B. A. Duar col., 0 4 November 1994. Distrito Federal, Po��o Azul (15 �� 34 ��� 36 ������ S, 48 ��02��� 30 ������ W, 1090m als), CHUNB 40854���40855, adult males, R. A. Brand��o col., 0 5 October 1994. CHUNB 14369���14376, juveniles, R. A. Brand��o col., 23 October 1994. Distrito Federal, Fazenda ��gua Limpa (15 �� 56 ��� 44 ������ S, 47 �� 54 ��� 34 ������ W, 1085m asl), CHUNB 25080, adult male, unknown collector, 23 November 2002. CHUNB 49652, adult male, P. H. Valdujo col., 14 February 2007. CHUNB 5079550796, adult males, G. R. Colli col., 27 November 2007. State of Goi��s, Cristalina Municipality (16 �� 44 ��� 30 ������ S, 47 �� 41 ��� 36 ������ W, 1050m asl), CHUNB 8369, adult male, G. R. Colli col., no date. CHUNB 14387, adult male, R. A. Brand��o, col., 22 May 1998. State of Goi��s, Piren��polis Municipality (15 �� 49 ���08������ S, 48 �� 59 ���08������ W, 880m asl), CHUNB 14386, adult male, R. A. Brand��o col., 22 June 1998. CHUNB 14380, adult male, R. A. Brand��o col., 0 1 September 1992. Reserva Particular do Patrim��nio Natural Vaga Fogo (15 �� 49 ��� 17 ������ S, 48 �� 59 ��� 44 ������ W, 810m asl), CHUNB 14390, adult male, M. G. Zatz col., 0 3 January 1999. Cocalzinho Municipality, Parque Estadual dos Pireneus (15 �� 48 ��� 16 ������ S, 48 �� 50 ���05������ W, 1270m asl), CHUNB 14388, adult male, R. A. Brand��o col., 28 December 1998. Novo Gama Municipality (16 ��03��� 24 ������ S, 48 ��01��� 21 ������ W, 1080m asl), CHUNB 14381, adult male, A. Sebben col., 10 November 1987. Diagnosis. The new species is diagnosed by the following combination of features: Medium size for the Bokermannohyla pseudopseudis group; dorsal coloration brownish ochre with darker scattered blotches; large eyes, forelimbs robust and short, with hypertrophied forearms in males; orange to reddish iris (bronze in some individuals); snout rounded in lateral and dorsal views (slightly acuminated in lateral view in some individuals); poorly-developed tibial fold, call frequency ranging from 0.5 to 0.7 kHz. Comparison with other species of the Bokermannohyla pseudopseudis species group. Bokermannohyla sapiranga differs from B. alvarengai (in parentheses) by having: 1) snout rounded in dorsal and lateral views (truncate snout in dorsal view and vertical in lateral view); 2) head slightly longer than wide (head wider than long); 3) tympanic fold present (absent); 4) tibial fold less conspicuous, not extending from the metacarpal external tubercle to the elbow (very conspicuous tibial fold, from the elbow to the basis of the external metacarpal tubercle); 5) dorsum with several scattered blotches (dorsum without pattern or large dark patches); 6) dorsum blotches extending to the inguinal region (inguinal region without blotches or pattern); 7) thigh bars poorly defined (conspicuous); 8) hidden surfaces of thighs with bars (hidden surfaces thighs without blotches, bars or patterns); 9) vent tubercles small, inconspicuous, and white (vent tubercles conspicuous); 10) gular region with small circular and darker blotches (gular region immaculate). Bokermannohyla sapiranga differs from B. flavopicta (in parentheses) by having: 1) dorsal background brownish with dark brown blotches (dorsal pattern lichenous-like, composed of small dark brown blotches in a light gray to light brown background); 2) similar head width and head length (heads wider than longer); 3) glandular tissues on mental region not evident (evident glandular tissues on mental region); 4) absence of yellow dots on the body in live individuals (small yellow dots present on lips, eyelids, loreal and gular regions, supratympanic fold, fore and hind limbs, flanks, and anal flap in live individuals); 5) smaller females [39.8���54.5 (N= 14) in B. sapiranga; 60.2���61.6 (N= 2) in B. flavopicta]. The new species differs from B. ibitiguara (in parentheses) by having: 1) snout rounded in dorsal view (truncate in dorsal view); 2) presence of several darker blotches on the gular region, that sometimes reach the chest (gular region immaculate white); 3) bars on legs poorly defined, becoming paler on hidden surfaces of thighs, and resembling diffuse blotches; 4) same pattern on the flanks and inguinal regions (transversal bars conspicuous, narrower, and regularly spaced in legs, flanks, and inguinal region); 5) dorsal coloration brown, with regular and darker blotches (dorsal color tan without blotches); 6) males with conspicuously hypertrophied forearms (males with forearm less hypertrophied). Bokermannohyla sapiranga differs from B. itapoty (in parentheses) by having: 1) tarsal fold poorly developed, from the metacarpal external tubercle to 4 / 5 of forearm length (tarsal fold more evident, from the metacarpal external tubercle to the elbow); 2) snout rounded in lateral and dorsal views (snout truncate in lateral and dorsal views); 3) tympanum oval-shaped (tympanum round); 4) absence of white spots on the dorsum, canthus rostralis, and loreal region (presence of white spots in some individuals); 5) presence of blotches on the hidden surface of thighs (thighs dark brown, without any bars or spots); 6) evident hypertrophied forearms in males (discrete hypertrophy in forearms of males). Bokermannohyla sapiranga differs from B. oxente (in parentheses) by having: 1) forearms of adult males more robust and hypertrophied (forearm width 3.65 �� 0.35, range: 3.08���3.97, N= 9 for B. oxente and 5.83 �� 0.70, range 5.02���7.68, N= 13 for B. sapiranga); 2) large dark blotches on dorsum (not present); 3) developed prepollex (less developed prepollex); 4) hidden surface of thighs with blotches (hidden thigh surface brownish, without blotches); 5) cloacal flap without white supracloacal stripe (white supracloacal stripe present). Bokermannohyla sapiranga can be distinguished from B. pseudopseudis (in parentheses) by having: 1) rounded snout in dorsal view (snout truncate in dorsal view) and rounded or slightly acuminated in lateral view (vertical); 2) iris red, orange, or bronze (iris yellowish or golden); 3) less conspicuous tarsal fold (more conspicuous); 4) dorsal surface of the fingers and toes discs grayish brown (whitish). Bokermannohyla sapiranga differs from B. sagarana (in parentheses) by having: 1) snout profile truncate in dorsal and lateral views (rounded in dorsal view, truncate in lateral view); 2) dorsal background brownish with dark brown blotches (lichenous-like dorsal color pattern: light gray with darker and lighter blotches of irregular shape); 3) bars on dorsal thigh surface brownish, poorly defined (dark gray perpendicular bars covering dorsal surface of thighs, highly contrasted with the background color). Bokermannohyla sapiranga differs from B. saxicola (in parentheses) by having: 1) snout profile rounded in dorsal and lateral view (rounded in dorsal view, truncate in lateral view); 2) brownish dorsum (dorsum light gray with darker blotches); 3) bars on thighs reaching the hidden outer surface, but poorly defined (bars on hidden thigh surface narrow, well defined); 4) head slender (head robust); conspicuous arm hypertrophy in males (discrete hypertrophy in males forearm); 5) absence of light stripe on the cloacal flap (light stripe on cloacal flap present); 6) forelimbs robust (forelimbs width: 5.83 �� 0.70, range 5.02���7.68, N= 17 in B. sapiranga; forelimbs width 3.78 �� 0.51, range 2.96���4.60, N= 7 in B. saxicola); 8) prepollex well developed (prepollex less developed). Description of holotype. Body robust (Fig. 1 and 2 A); head slightly longer than wider; snout rounded in lateral (Fig. 2 B) and in dorsal views; nostrils slightly protuberant, nostril aperture oval; eyes large, slightly protruding laterally; tympanum large, rounded; maximum eye length 1.5 times the maximum tympanum length; supratympanic fold evident, well-developed, extending from the posterior eye margin to the shoulder, at the level of the inferior tympanum margin. Vocal sac not externally expanded; vocal slits present; tongue large, wider than longer, almost cordiform; vomerine teeth disposed in two continuous rows, placed posterior to the choanae, forming an open angle (ten on the right and 9 on the left side); choanae large, almost circular, apart from each other; hypertrophied forearms (Fig. 1 and Fig. 2 B���C); external forearm fold evident, extending from the elbow to the wrist; well-developed prepollex, with a sharp spine, not exposed; slender fingers; subarticular tubercles simple, round, and well-developed; few round and discrete palmar tubercles; a large external metacarpal tubercle, fanshaped, partially divided in three parts (united at the basis) by two grooves on the internal margin; finger pads ovoid, single, well-developed; relative finger lengths IV>II>III>V; webbing hand formula II���III 2 -��� 3 - IV 3 ��� 2 +V (Fig. 2 C); nuptial excrescence small, discrete, round, placed on the dorsal surface of finger II basis, at the prepollex junction; legs slender, thigh longer than tibia (Fig. 2 D); tarsal folds evident; internal tarsal fold from the posterior margin of the internal metatarsal tubercle to the heel (Fig. 2 D); external metatarsal fold less developed, extending two thirds the tarsal length from the first phalanx of toe V; internal metatarsal tubercle well developed, ovoid; external metatarsal tubercle small and round; subarticular tubercles simple and round; plantar tubercles small, discrete, round; relative toe length IV>V>III>II>I; web foot formula I 1 +��� 2 II 1 +��� 1 III 1 +��� 2 IV 2 ��� 1 V (Fig. 2 E). Toe discs slightly less developed than finger discs; dorsal texture moderately granular; ventral region, chest, lateral flanks, and ventral thigh surface densely granular (Fig. 2 D); ventral tibia and ventral mandible surfaces smooth; vent region dark; an evident fold delimits the dorsum and the cloacal flap. Measurements of holotype (in mm). SVL: 46.98, HL: 16.25, HW: 16.86, ED: 5.51, IOD: 7.86, END: 4.46, IND: 4.53, NSD: 3.28, TD: 2.95, PPL: 4.68, FDD: 1.90, TGL: 22.91, TBL: 25.52, FTL: 23.47, TDD: 1.83, FL: 7.48, FW: 5.75, HD: 15.72, AL: 6.56, TAL: 12.93. Color in preservative. Dorsum brownish gray, with large dark brown or gray blotches; large dorsal dark brown blotches similar in size; flanks light gray with large dark brown or gray blotches; venter immaculate, whitish; gular region with several pale, brownish blotches; ventral tarsal surface brownish gray; dorsal surface of limbs brownish gray; ventral surfaces of toe discs white; hidden surfaces of the thighs dark brown. Color in life. Dorsum and limbs brownish, with large darker brown blotches; iris reddish (Fig. 1 A), tympanum and foot webbing brownish; venter light ochre. Morphometric analyses. Measurements of Bokermannohyla sapiranga are shown in Table 1. To evaluate morphometric differences between B. sapiranga and B. pseudopseudis we used the model selection with shape variables. In this case, eight models were selected, all combining more than nine variables (Table 2). Just two models presented AICw value greater than 15.0. However, all the models selected share several variables. The variables snout-vent length, eye diameter, and tibia length were less important to explain the average model (Table 3). These variables were not recovered in any of the eight models selected by the AIC criterion (Table 2). The average model explained less than half of the differences between B. pseudopseudis and B. sapiranga (r �� = 0.413; r �� adjusted = 0.291). The variables head length, interorbital distance, head width, and fourth toe disc diameter were the four most important selected by the average model (Table 3), suggesting that the differences between B. pseudopseudis and the new species were related to the head proportions and size of the toe discs. The PCA suggests that Bokermannohyla pseudopseudis and B. sapiranga are slightly different on general morphology (Fig. 3). The first two scores of the PCA account for 61.34 % of the variance. The first component accounts for 40.48 % of the variance and showed higher positive values for nostril-snout distance and interorbital distance, and higher negative values for third finger disc diameter, and fourth toe disc diameter (Table 4). The second component accounts for 20.86 % of the variance, presenting higher positive value for third finger disc diameter, and higher negative value for tympanum diameter. Advertisement call. The advertisement call of Bokermannohyla sapiranga from the type locality (Bras��lia municipality) consists of series of 5 to 7 notes with no harmonic structure (Fig. 4). Some notes show a pulsed structure (Fig. 4 A). Mean call duration was 997 �� 16.4 ms (range = 759���1202 ms, N= 5, Fig. 4 A). Mean note duration was 98.4 �� 9.3 ms (range = 85���116 ms), and mean internote interval was 110 �� 16.5 ms (range = 92���132 ms, Fig. 4 A). Dominant frequency did not vary (645.9 Hz, Fig. 4 B). Temporal call parameters from Bras��lia were slightly longer when compared to calls from Piren��polis, in number of notes (3���5), mean call duration 520 �� 14.2 ms (range = 380���790 ms) (N= 13) and duration of notes (mean note duration = 67.5 �� 6.2 ms, ranging 75���85 ms). Mean dominant frequency was also slightly shorter in Piren��polis 559.9 �� 49.7 Hz (range = 516.8���602.9 Hz). The call of B. sapiranga is similar to calls of other species in the B. pseudopseudis group (Table 5). It is more similar to B. pseudopseudis, but readily separated from it by lacking the harmonic structure showed by B. pseudopseudis (pulsionated in B. sapiranga), along with differences in spectral features, number and duration of notes, and call length (Table 5). The maximum frequency in B. sapiranga is under 1.0 kHz, whereas in B. pseudopseudis it may reach 2.4 kHz (Eterovick & Brand��o, 2001; Lugli & Haddad, 2006 b). Mean call duration of B. pseudopseudis was 2.6 s (Eterovick & Brand��o 2001), while ranging 0.4��� 1.2 s in B. sapiranga. pure notes, without B. pseudopseudis 3 5���10 64 ���84 56.4���71.8 0.4���2.4 up to five visible pulses most notes are B. sapiranga 4 5���7 85 ���116 92���132 0.5���0.7 not present pulsed Three main pure notes, without B. saxicola 5 25 ��� 25 22 ���441 602 0.9���4.5 harmonics visible pulses Source for comparisons: 1 Cardoso (1983); 2 Lugli & Haddad (2006 b); 3 Eterovick & Brand��o (2001); 4 Present work (call parameters based in the specimen recorded from Bras��lia municipality, type locality); 5 Bokermann (1964) and Eterovick & Brand��o (2001). Etymology. In Tupi indigenous language, sapiranga means red eye, an allusion to the reddish iris in the most individuals of the species. The specific epipeth is also homage to Marco (Sapiranga) Freitas, for his continuous efforts to popularize the Brazilian herpetofauna. Distribution. Bokermannohyla sapiranga is known from the type locality (Bras��lia, Distrito Federal), and the Municipalities of Cristalina, Novo Gama, Catal��o, and Piren��polis, State of Goi��s (Fig. 5). Ecology and natural history notes. Bokermannohyla sapiranga and B. pseudopseudis are species associated with streams and rivulets with fast water flow in highlands of Central Brazil. Basking behavior was observed for both species, in Bras��lia (B. sapiranga) (Fig. 6 A) and Parque Nacional da Chapada dos Veadeiros (B. pseudopseudis). However, they present some ecological differences. Bokermannohyla pseudopseudis uses streams in open areas and rivulets with rocky beds, while B. sapiranga besides using similar habitats in some localities (e.g. Piren��polis, Cristalina, and Bras��lia Municipalities) also occurs in dense wet gallery forest, calling near small waterfalls formed by roots and logs along rivulet (Bras��lia Municipality). In these habitats, tadpoles are found in pools and backwaters with mud beds. We never found B. pseudopseudis using such kind of habitat. As other species of the B. pseudopseudis group, B. sapiranga shows low ability to perch on the vegetation. Most individuals were observed on rocks, soil, or dead logs. In Bras��lia Municipality, Ameerega flavopicta, Aplastodiscus aff. perviridis, Barycholos ternetzi, Hypsiboas lundii, Hypsiboas goianus, Rhinella cerradensis, Rhinella rubescens, and Scinax skaios can be found in the same habitats where B. sapiranga occurs., Published as part of Brand��o, Reuber Albuquerque, Magalh��es, Rafael F��lix De, Garda, Adrian Antonio, Campos, Leandro Abr��sio, Sebben, Antonio & Maciel, Natan Medeiros, 2012, A new species of Bokermannohyla (Anura: Hylidae) from highlands of Central Brazil, pp. 28-42 in Zootaxa 3527 on pages 30-38, DOI: 10.5281/zenodo.213546, {"references":["Eterovick, P. C. & Brandao, R. A. (2001) A description of the tadpoles and advertisement calls of members of the Hyla pseudopseudis group. Journal of Herpetology, 35, 442 - 450.","Lugli, L. & Haddad, C. F. B. (2006 b) A new species of the Bokermannohyla pseudopseudis group from Central Bahia, Brazil (Amphibia, Hylidae). Herpetologica, 62, 453 - 465.","Cardoso, A. J. (1983) Descricao e biologia de uma nova especie de Hyla Laurenti, 1768 (Amphibia, Anura, Hylidae). Iheringia, Serie Zoologia, 62, 37 - 45.","Bokermann, W. C. A. (1964) Dos nuevas especies de Hyla de Minas Gerais y notas sobre Hyla alvarengai Bok. (Amphibia, Salientia, Hylidae). Neotropica, 10, 67 - 76."]}

Bokermannohyla sapiranga sp. nov. (Figs. 1–2) Holotype. Brazil, Distrito Federal, Reserva Ecológica do Roncador (15 ° 55 ’ 49 ” S, 47 ° 52 ’ 59 ” W, 1110m asl), CHUNB 62384, adult male, R.A. Brandão, R.D. Françoso, T. Marques, R.M. Japiassu, and L.R.A. Braga col., 10 December 2009. Paratypes. Brazil, Distrito Federal, Brasília (15 ° 52 ’ 25 ” S, 47 ° 51 ’ 59 ” W, 1070m asl), CFBH 2248–2249, adult males, A. Sebben col., no date. CHUNB 14383–14384, adult females, A. Sebben col., 0 1 April 1986. CHUNB 38832, adult male, unknown collector, 0 1 May 1993. CHUNB 14377–14379, juveniles, B. A. Duar col., 0 4 November 1994. Distrito Federal, Poço Azul (15 ° 34 ’ 36 ’’ S, 48 °02’ 30 ’’ W, 1090m als), CHUNB 40854–40855, adult males, R. A. Brandão col., 0 5 October 1994. CHUNB 14369–14376, juveniles, R. A. Brandão col., 23 October 1994. Distrito Federal, Fazenda Água Limpa (15 ° 56 ’ 44 ’’ S, 47 ° 54 ’ 34 ’’ W, 1085m asl), CHUNB 25080, adult male, unknown collector, 23 November 2002. CHUNB 49652, adult male, P. H. Valdujo col., 14 February 2007. CHUNB 5079550796, adult males, G. R. Colli col., 27 November 2007. State of Goiás, Cristalina Municipality (16 ° 44 ’ 30 ’’ S, 47 ° 41 ’ 36 ’’ W, 1050m asl), CHUNB 8369, adult male, G. R. Colli col., no date. CHUNB 14387, adult male, R. A. Brandão, col., 22 May 1998. State of Goiás, Pirenópolis Municipality (15 ° 49 ’08’’ S, 48 ° 59 ’08’’ W, 880m asl), CHUNB 14386, adult male, R. A. Brandão col., 22 June 1998. CHUNB 14380, adult male, R. A. Brandão col., 0 1 September 1992. Reserva Particular do Patrimônio Natural Vaga Fogo (15 ° 49 ’ 17 ’’ S, 48 ° 59 ’ 44 ’’ W, 810m asl), CHUNB 14390, adult male, M. G. Zatz col., 0 3 January 1999. Cocalzinho Municipality, Parque Estadual dos Pireneus (15 ° 48 ’ 16 ’’ S, 48 ° 50 ’05’’ W, 1270m asl), CHUNB 14388, adult male, R. A. Brandão col., 28 December 1998. Novo Gama Municipality (16 °03’ 24 ’’ S, 48 °01’ 21 ’’ W, 1080m asl), CHUNB 14381, adult male, A. Sebben col., 10 November 1987. Diagnosis. The new species is diagnosed by the following combination of features: Medium size for the Bokermannohyla pseudopseudis group; dorsal coloration brownish ochre with darker scattered blotches; large eyes, forelimbs robust and short, with hypertrophied forearms in males; orange to reddish iris (bronze in some individuals); snout rounded in lateral and dorsal views (slightly acuminated in lateral view in some individuals); poorly-developed tibial fold, call frequency ranging from 0.5 to 0.7 kHz. Comparison with other species of the Bokermannohyla pseudopseudis species group. Bokermannohyla sapiranga differs from B. alvarengai (in parentheses) by having: 1) snout rounded in dorsal and lateral views (truncate snout in dorsal view and vertical in lateral view); 2) head slightly longer than wide (head wider than long); 3) tympanic fold present (absent); 4) tibial fold less conspicuous, not extending from the metacarpal external tubercle to the elbow (very conspicuous tibial fold, from the elbow to the basis of the external metacarpal tubercle); 5) dorsum with several scattered blotches (dorsum without pattern or large dark patches); 6) dorsum blotches extending to the inguinal region (inguinal region without blotches or pattern); 7) thigh bars poorly defined (conspicuous); 8) hidden surfaces of thighs with bars (hidden surfaces thighs without blotches, bars or patterns); 9) vent tubercles small, inconspicuous, and white (vent tubercles conspicuous); 10) gular region with small circular and darker blotches (gular region immaculate). Bokermannohyla sapiranga differs from B. flavopicta (in parentheses) by having: 1) dorsal background brownish with dark brown blotches (dorsal pattern lichenous-like, composed of small dark brown blotches in a light gray to light brown background); 2) similar head width and head length (heads wider than longer); 3) glandular tissues on mental region not evident (evident glandular tissues on mental region); 4) absence of yellow dots on the body in live individuals (small yellow dots present on lips, eyelids, loreal and gular regions, supratympanic fold, fore and hind limbs, flanks, and anal flap in live individuals); 5) smaller females [39.8–54.5 (N= 14) in B. sapiranga; 60.2–61.6 (N= 2) in B. flavopicta]. The new species differs from B. ibitiguara (in parentheses) by having: 1) snout rounded in dorsal view (truncate in dorsal view); 2) presence of several darker blotches on the gular region, that sometimes reach the chest (gular region immaculate white); 3) bars on legs poorly defined, becoming paler on hidden surfaces of thighs, and resembling diffuse blotches; 4) same pattern on the flanks and inguinal regions (transversal bars conspicuous, narrower, and regularly spaced in legs, flanks, and inguinal region); 5) dorsal coloration brown, with regular and darker blotches (dorsal color tan without blotches); 6) males with conspicuously hypertrophied forearms (males with forearm less hypertrophied). Bokermannohyla sapiranga differs from B. itapoty (in parentheses) by having: 1) tarsal fold poorly developed, from the metacarpal external tubercle to 4 / 5 of forearm length (tarsal fold more evident, from the metacarpal external tubercle to the elbow); 2) snout rounded in lateral and dorsal views (snout truncate in lateral and dorsal views); 3) tympanum oval-shaped (tympanum round); 4) absence of white spots on the dorsum, canthus rostralis, and loreal region (presence of white spots in some individuals); 5) presence of blotches on the hidden surface of thighs (thighs dark brown, without any bars or spots); 6) evident hypertrophied forearms in males (discrete hypertrophy in forearms of males). Bokermannohyla sapiranga differs from B. oxente (in parentheses) by having: 1) forearms of adult males more robust and hypertrophied (forearm width 3.65 ± 0.35, range: 3.08–3.97, N= 9 for B. oxente and 5.83 ± 0.70, range 5.02–7.68, N= 13 for B. sapiranga); 2) large dark blotches on dorsum (not present); 3) developed prepollex (less developed prepollex); 4) hidden surface of thighs with blotches (hidden thigh surface brownish, without blotches); 5) cloacal flap without white supracloacal stripe (white supracloacal stripe present). Bokermannohyla sapiranga can be distinguished from B. pseudopseudis (in parentheses) by having: 1) rounded snout in dorsal view (snout truncate in dorsal view) and rounded or slightly acuminated in lateral view (vertical); 2) iris red, orange, or bronze (iris yellowish or golden); 3) less conspicuous tarsal fold (more conspicuous); 4) dorsal surface of the fingers and toes discs grayish brown (whitish). Bokermannohyla sapiranga differs from B. sagarana (in parentheses) by having: 1) snout profile truncate in dorsal and lateral views (rounded in dorsal view, truncate in lateral view); 2) dorsal background brownish with dark brown blotches (lichenous-like dorsal color pattern: light gray with darker and lighter blotches of irregular shape); 3) bars on dorsal thigh surface brownish, poorly defined (dark gray perpendicular bars covering dorsal surface of thighs, highly contrasted with the background color). Bokermannohyla sapiranga differs from B. saxicola (in parentheses) by having: 1) snout profile rounded in dorsal and lateral view (rounded in dorsal view, truncate in lateral view); 2) brownish dorsum (dorsum light gray with darker blotches); 3) bars on thighs reaching the hidden outer surface, but poorly defined (bars on hidden thigh surface narrow, well defined); 4) head slender (head robust); conspicuous arm hypertrophy in males (discrete hypertrophy in males forearm); 5) absence of light stripe on the cloacal flap (light stripe on cloacal flap present); 6) forelimbs robust (forelimbs width: 5.83 ± 0.70, range 5.02–7.68, N= 17 in B. sapiranga; forelimbs width 3.78 ± 0.51, range 2.96–4.60, N= 7 in B. saxicola); 8) prepollex well developed (prepollex less developed). Description of holotype. Body robust (Fig. 1 and 2 A); head slightly longer than wider; snout rounded in lateral (Fig. 2 B) and in dorsal views; nostrils slightly protuberant, nostril aperture oval; eyes large, slightly protruding laterally; tympanum large, rounded; maximum eye length 1.5 times the maximum tympanum length; supratympanic fold evident, well-developed, extending from the posterior eye margin to the shoulder, at the level of the inferior tympanum margin. Vocal sac not externally expanded; vocal slits present; tongue large, wider than longer, almost cordiform; vomerine teeth disposed in two continuous rows, placed posterior to the choanae, forming an open angle (ten on the right and 9 on the left side); choanae large, almost circular, apart from each other; hypertrophied forearms (Fig. 1 and Fig. 2 B–C); external forearm fold evident, extending from the elbow to the wrist; well-developed prepollex, with a sharp spine, not exposed; slender fingers; subarticular tubercles simple, round, and well-developed; few round and discrete palmar tubercles; a large external metacarpal tubercle, fanshaped, partially divided in three parts (united at the basis) by two grooves on the internal margin; finger pads ovoid, single, well-developed; relative finger lengths IV>II>III>V; webbing hand formula II–III 2 -– 3 - IV 3 – 2 +V (Fig. 2 C); nuptial excrescence small, discrete, round, placed on the dorsal surface of finger II basis, at the prepollex junction; legs slender, thigh longer than tibia (Fig. 2 D); tarsal folds evident; internal tarsal fold from the posterior margin of the internal metatarsal tubercle to the heel (Fig. 2 D); external metatarsal fold less developed, extending two thirds the tarsal length from the first phalanx of toe V; internal metatarsal tubercle well developed, ovoid; external metatarsal tubercle small and round; subarticular tubercles simple and round; plantar tubercles small, discrete, round; relative toe length IV>V>III>II>I; web foot formula I 1 +– 2 II 1 +– 1 III 1 +– 2 IV 2 – 1 V (Fig. 2 E). Toe discs slightly less developed than finger discs; dorsal texture moderately granular; ventral region, chest, lateral flanks, and ventral thigh surface densely granular (Fig. 2 D); ventral tibia and ventral mandible surfaces smooth; vent region dark; an evident fold delimits the dorsum and the cloacal flap. Measurements of holotype (in mm). SVL: 46.98, HL: 16.25, HW: 16.86, ED: 5.51, IOD: 7.86, END: 4.46, IND: 4.53, NSD: 3.28, TD: 2.95, PPL: 4.68, FDD: 1.90, TGL: 22.91, TBL: 25.52, FTL: 23.47, TDD: 1.83, FL: 7.48, FW: 5.75, HD: 15.72, AL: 6.56, TAL: 12.93. Color in preservative. Dorsum brownish gray, with large dark brown or gray blotches; large dorsal dark brown blotches similar in size; flanks light gray with large dark brown or gray blotches; venter immaculate, whitish; gular region with several pale, brownish blotches; ventral tarsal surface brownish gray; dorsal surface of limbs brownish gray; ventral surfaces of toe discs white; hidden surfaces of the thighs dark brown. Color in life. Dorsum and limbs brownish, with large darker brown blotches; iris reddish (Fig. 1 A), tympanum and foot webbing brownish; venter light ochre. Morphometric analyses. Measurements of Bokermannohyla sapiranga are shown in Table 1. To evaluate morphometric differences between B. sapiranga and B. pseudopseudis we used the model selection with shape variables. In this case, eight models were selected, all combining more than nine variables (Table 2). Just two models presented AICw value greater than 15.0. However, all the models selected share several variables. The variables snout-vent length, eye diameter, and tibia length were less important to explain the average model (Table 3). These variables were not recovered in any of the eight models selected by the AIC criterion (Table 2). The average model explained less than half of the differences between B. pseudopseudis and B. sapiranga (r ² = 0.413; r ² adjusted = 0.291). The variables head length, interorbital distance, head width, and fourth toe disc diameter were the four most important selected by the average model (Table 3), suggesting that the differences between B. pseudopseudis and the new species were related to the head proportions and size of the toe discs. The PCA suggests that Bokermannohyla pseudopseudis and B. sapiranga are slightly different on general morphology (Fig. 3). The first two scores of the PCA account for 61.34 % of the variance. The first component accounts for 40.48 % of the variance and showed higher positive values for nostril-snout distance and interorbital distance, and higher negative values for third finger disc diameter, and fourth toe disc diameter (Table 4). The second component accounts for 20.86 % of the variance, presenting higher positive value for third finger disc diameter, and higher negative value for tympanum diameter. Advertisement call. The advertisement call of Bokermannohyla sapiranga from the type locality (Brasília municipality) consists of series of 5 to 7 notes with no harmonic structure (Fig. 4). Some notes show a pulsed structure (Fig. 4 A). Mean call duration was 997 ± 16.4 ms (range = 759–1202 ms, N= 5, Fig. 4 A). Mean note duration was 98.4 ± 9.3 ms (range = 85–116 ms), and mean internote interval was 110 ± 16.5 ms (range = 92–132 ms, Fig. 4 A). Dominant frequency did not vary (645.9 Hz, Fig. 4 B). Temporal call parameters from Brasília were slightly longer when compared to calls from Pirenópolis, in number of notes (3–5), mean call duration 520 ± 14.2 ms (range = 380–790 ms) (N= 13) and duration of notes (mean note duration = 67.5 ± 6.2 ms, ranging 75–85 ms). Mean dominant frequency was also slightly shorter in Pirenópolis 559.9 ± 49.7 Hz (range = 516.8–602.9 Hz). The call of B. sapiranga is similar to calls of other species in the B. pseudopseudis group (Table 5). It is more similar to B. pseudopseudis, but readily separated from it by lacking the harmonic structure showed by B. pseudopseudis (pulsionated in B. sapiranga), along with differences in spectral features, number and duration of notes, and call length (Table 5). The maximum frequency in B. sapiranga is under 1.0 kHz, whereas in B. pseudopseudis it may reach 2.4 kHz (Eterovick & Brandão, 2001; Lugli & Haddad, 2006 b). Mean call duration of B. pseudopseudis was 2.6 s (Eterovick & Brandão 2001), while ranging 0.4– 1.2 s in B. sapiranga. pure notes, without B. pseudopseudis 3 5–10 64 –84 56.4–71.8 0.4–2.4 up to five visible pulses most notes are B. sapiranga 4 5–7 85 –116 92–132 0.5–0.7 not present pulsed Three main pure notes, without B. saxicola 5 25 – 25 22 –441 602 0.9–4.5 harmonics visible pulses Source for comparisons: 1 Cardoso (1983); 2 Lugli & Haddad (2006 b); 3 Eterovick & Brandão (2001); 4 Present work (call parameters based in the specimen recorded from Brasília municipality, type locality); 5 Bokermann (1964) and Eterovick & Brandão (2001). Etymology. In Tupi indigenous language, sapiranga means red eye, an allusion to the reddish iris in the most individuals of the species. The specific epipeth is also homage to Marco (Sapiranga) Freitas, for his continuous efforts to popularize the Brazilian herpetofauna. Distribution. Bokermannohyla sapiranga is known from the type locality (Brasília, Distrito Federal), and the Municipalities of Cristalina, Novo Gama, Catalão, and Pirenópolis, State of Goiás (Fig. 5). Ecology and natural history notes. Bokermannohyla sapiranga and B. pseudopseudis are species associated with streams and rivulets with fast water flow in highlands of Central Brazil. Basking behavior was observed for both species, in Brasília (B. sapiranga) (Fig. 6 A) and Parque Nacional da Chapada dos Veadeiros (B. pseudopseudis). However, they present some ecological differences. Bokermannohyla pseudopseudis uses streams in open areas and rivulets with rocky beds, while B. sapiranga besides using similar habitats in some localities (e.g. Pirenópolis, Cristalina, and Brasília Municipalities) also occurs in dense wet gallery forest, calling near small waterfalls formed by roots and logs along rivulet (Brasília Municipality). In these habitats, tadpoles are found in pools and backwaters with mud beds. We never found B. pseudopseudis using such kind of habitat. As other species of the B. pseudopseudis group, B. sapiranga shows low ability to perch on the vegetation. Most individuals were observed on rocks, soil, or dead logs. In Brasília Municipality, Ameerega flavopicta, Aplastodiscus aff. perviridis, Barycholos ternetzi, Hypsiboas lundii, Hypsiboas goianus, Rhinella cerradensis, Rhinella rubescens, and Scinax skaios can be found in the same habitats where B. sapiranga occurs.

Rhinella cerradensis sp. nov. (Figs. 1–3) Holotype: Coleção Herpetológica da Universidade de Brasília (CHUNB 49573), adult male collected at Centro de Instrução e Adestramento de Brasília, Brasília, Distrito Federal, Brazil (14 ° 49 'S; 45 ° 58 'W) in November 2002, by N.M. Maciel, R.H. Matsushita, A.Q.Teixeira Jr., and B. A. Duar. Paratypes: CHUNB 40276, 40277, 40280, 49281 (males), 40278, 40279 (females), collected along with the holotype; CHUNB 40273–40275 (females), collected at Rio Pratudão Farm, Correntina Municipality, Bahia State, in October 1996 by R.A. Brandão; CHUNB 13790–13791 (females), collected at Emas National Park, Mineiros Municipality, Goiás State, in November 1998 by R.A. Brandão; CHUNB 38669 (male), collected at Jardim Botânico, Brasília, Distrito Federal, in January 2001 by B.A. Duar; CHUNB 25043 (female), collected at Brasília, Distrito Federal, in September 2001 by O. R. Pires Jr.; CHUNB 38655, 38656 (females), 38658 (juvenile), collected at Trijunção Farm, Cocos Municipality, Bahia State, in October 2001, by R.A. Brandão; CHUNB 29386 (male), collected at Brasília, Distrito Federal, in August 2002 by D.S. Diniz; CHUNB 39979–39980 (females), collected at Trijunção Farm, Cocos Municipality, Bahia State, (data not recorded) by R.A. Brandão; CHUNB 39953 (female), collected at Centro de Instrução e Adestramento de Brasília, Brasília, Distrito Federal, in January 2003, by N.M. Maciel; CHUNB 38668 (male), collected at Centro de Instrução e Adestramento de Brasília, Brasília, Distrito Federal, in August, 2003, by N.M. Maciel, R.H. Matsushita, and R.A. Brandão (advertisement call recorded); CHUNB 38667 (female), 38670–38671 (males), collected at Trijunção Farm, Cocos Municipality, Bahia State, in October 2004, by R.A. Brandão, N.M. Maciel and A. Sebben; CHUNB 43864 (female), collected at São Domingos Municipality, Goiás State, in November 2004. Species measurements are provided in Table 1. Diagnosis and comparisons with other species: A medium to large sized species, similar to R. icterica, R. marina, R. poeppigii, R. schneideri, and R. jimi mostly by the well developed cranial crests, long and broad parotoid gland, head wider than long, and morphometric features (see morphometric analysis section). Rhinella cerradensis has the following unique combination of features within the R. marina group: medium to large size, (males 120.4 (± 13.4) mm, ranging 103.3–147.1 mm and females 114.5 (± 20.1) mm, ranging 82.1– 139.8 mm); head wider than long; well developed cranial crests, with evident spicules; snout in dorsal view mucronate to truncate and obtuse in lateral view; a medium sized parotoid gland, parotoid gland elliptical, yellowish, with a high density of keratinous pointed spicules; pointed spicules present on dorsal glands, fore, and hind limbs; tibial gland absent; dorsum uniform brownish to dark green and venter brownish white in males; females with brownish green dorsum, but with black coloration in many areas (sometimes also in hind limbs), resembling R. icterica female coloration; a jagged or “scalloped” suture formed by the articulation between the medial ramus of the pterygoid and the parasphenoid alae. The new species can be quickly distinguished from R. schneideri and R. jimi by the smaller body and parotoid gland size and by the absent of tibial gland (PGL in R. schneideri 28.9 (± 9.8) mm (n= 27), R. jimi 34.8 (± 11.8) mm (n= 9), and R. cerradensis 27.5 (± 7.7) mm (n= 18)). From R. icterica the new species can be distinguished by size of parotoid gland (larger and rounded in R. icterica, PGL = 29.7 (± 9.6) mm (n= 29)); by a less developed preocular crest; by narrow head (wide in R. icterica); by tympanum without folded skin (present in R. icterica); and by smaller hand (HAL in R. cerradensis 25.7 (±6.0) mm (n= 18) and in R. icterica 26.7 (± 6.1) mm (n= 29)). From R. marina, R. cerradensis can be distinguished by shape of parotoid gland (almost rounded in R. marina); by narrower and longer head (wider in R. marina). From R. veredas, it can be easily distinguished by complete supraocular crest and by dorsal coloration of the male. From R. rubescens, it can be distinguished by its larger size (SVL= 115.5 (± 26.4) mm (n= 18) in R. cerradensis and 69.1 (± 23.4) mm in R. rubescens (n= 49)); shape of parotoid gland (long and narrow in R. rubescens); wider head (LC= 23.2 (± 7.4) mm in R. rubescens (n= 49)); snout profile (sub-elliptical in dorsal view in R. rubescens), cranial crests developed (weakly developed in R. rubescens); and dorsal glands with keratinous spicules on body (smooth in R. rubescens). From R. arenarum, it can be distinguished by larger size (SVL= 87.3 (±11.0) mm in R. arenarum (n= 25)); parotoid gland shape (long and narrow in R. arenarum), by more developed cranial crests (weakly developed in R. arenarum); tongue long and narrow (rounded and short in R. arenarum); and by dorsal male coloration. From R. achavali it can be distinguished by larger size (SVL= 95.4 (± 6.2) mm in R. achavali (n= 5)), shape of parotoid glands (long and narrow in R. achavali); by dorsal male coloration; by pointed spiculae on body glands; and by the rough skin. Description of the holotype: General aspect robust (Fig. 1 A). Head wider than long; SVL approximately 2.7 × head width; snout mucronate to truncate in dorsal view and obtuse in lateral view (Figs. 3 A and 3 B); almost all cranial crests present and developed, parietal crest absent; pre-tympanic and rostral crests well developed; supranasal crest evident with rugosites, produced by cranial striations; supratympanic crest developed, in contact with the parotoid gland; cranial crests keratinized, especially the supraocular crest; loreal region smooth; nostrils large, with lateral elliptic openings directed upward; interorbital distance and eye diameter equal in length; eye diameter about 1.5 × tympanum diameter; eyelid large, with small keratinous spicules; tympanum visible, well developed, rounded; dorsum of head flat, with small rounded spicules. Parotoid gland evident, elliptical, with keratinous spicules; parotoid length about 2.6 × its width. Tongue longer than wide, smooth, with one single superficial central fold, free in its posterior end, attached to jaw; vomerine teeth absent; choanae elliptical. Fingers thin and developed (Fig. 3 C); forearm and arm robust; relative finger length II Morphometric analyses: The PCA suggests the presence of two morphological subgroups in species of R. marina group (Fig. 4). A subgroup of species represented by the white symbols, composed by R. marina, R. icterica, R. schneideri, R. jimi, and R. cerradensis and another represented by the black symbols, consisting of R. arenarum, R. rubescens, R. achavali, and R. veredas. The first two scores account for a total of 94.8 % of the variance. The first component accounts for 88.4 % of the variance and showed high and positive scores. The second component accounts for 6.4 % of the variance and showed positive higher values for tympanum diameter and foot and leg lengths, and negative values for parotoid gland length (Table 2). Examination of factor loadings suggested the formation of two subgroups. The first subgroup (white symbols) is composed of the larger sized species, with developed and marked cranial crests, relatively smaller tympanum, and shorter limb length. The second subgroup (black symbols) is formed by the smaller species with less developed cranial crests and relatively longer parotoid glands (Fig. 4). Coloration: Sexual dimorphism is well marked in coloration (Fig. 2). In life, males have a brownish green dorsum (sometimes a dark green dorsum) and a brownish white venter. Females have the same brownish green dorsum and brownish white venter of the male, but show black coloration in many areas of dorsum (sometimes also on hind limbs). In this respect, the females of this species resemble R. icterica females. The coloration of preserved specimens resembles the coloration of males and females in life, but the parotoid glands turn yellowish. Description of the tadpole: Descriptions are based on a tadpole series (CHUNB 49574) collected at Estação Rádio, Centro de Instrução e Adestramento de Brasília, Brasília Municipality, Distrito Federal on 31 August 2002, by N.M. Maciel and R.H. Matsushita. Tadpole series measurements are provided in Table 3. Measurements of the specimen used for description (Fig. 5) (in mm): BL 9.72; IND 1.30; IOD 2.90; MTH 4.30; TAL 12.64; TL 21.76; TMH 1.50; TMW 1.65. The specimen used for description is in stage 40 (Gosner, 1960) (Fig. 5). Body elliptical in lateral (Fig. 5 A) and dorsal views (Fig. 5 B). Body 76 % of tail length and 45 % of total length; body wider than deep; snout rounded in lateral and dorsal views; eyes dorsal, directed laterally; distance between eyes 2.2 × the distance between nostrils; nostril opening kidney shape; spiracle sinistral, near to midbody axis, opening in the body wall, without external tube. Vent tube medial (easily visible in tadpoles in early stages). Tail musculature gradually tapering to rounded tip. Caudal fin fusiform; dorsal fin about same height as ventral fin. Color in preservative uniform, dark brown. Oral disc anteroventral, and emarginated laterally in lower and upper jaw. LTRF 2 (2)/ 3 (1); A 2 slightly longer than A 1, P 1 slightly shorter than A 1 and longer than P 2 and P 3. Lower jaw serrate and shallowly V-shaped. The majority of tadpole specimens are in stages 40–42. The tadpoles in stage 42 differed from those in stage 40 mainly by the development of hindlimbs, presence of emerged forelimbs, more rounded snout, and shorter body. The tadpoles described from species of the R. marina group and those of the new species are similar in general shape (Rosa, 1951; Savage, 1960; Rosa, 1965; Cei, 1980; Heyer et al., 1990; Eterovick & Sazima, 1999). According to Eterovick & Sazima (1999) the tadpoles of R. rubescens are more similar to those of R. schneideri in body proportions and spiracle position, while the tadpoles of R. arenarum more closely resemble those of R. icterica. The tadpoles of the new species resemble R. rubescens and R. schneideri in body proportions and spiracle position. The tadpole of R. cerradensis can be diagnosed by body proportions, spiracle position, and the absence of an external spiracular tube, with the opening in the body wall (Fig. 5 A). The absence of an external tube is unique among the tadpoles described of the R. marina group. Osteological features: Skull with the robust general aspect of the species of Rhinella marina group (sensu Pramuk, 2006). The specimens analyzed have extremely well-ossified and exostosed crania that are ornamented with deep striations, pits, and rugosities. The cranium is wider than long, with the greatest width at the level of the quadratojugals, as it is characteristic of other species of the group. All cranial crests present, with the exception of the parietal crest (Fig. 6). The Arabic numbers between quotations marks in the text indicate skull features of the new species (see Fig. 6). A detailed description of cranial features follows. Alary process of premaxilla postero-lateraly positioned; premaxilary bones are miss in the figure, frequently split during osteological bufonid preparations; vertical process of maxilla flattened laterally but dilated medially, maxillary crest well developed (5) (Fig. 6 B and C); nasals in contact medially (1), anterior tips of nasals ventrally projected, ornamentations along edges forming the canthal (3) and supraorbital crests (4) (Fig. 6 A); vertical arm of nasals robust on top and flattened on bottom; anterior portion of frontoparietal with rectangular shape, posterior portion projected at posterior end of skull, composing apical part of foramen magnum (2); frontoparietal not contacting prootic process (2) (Fig. 6 A); medial area of frontoparietal large, with a portion that does not compound cranial box; this lateral portion bends in a plane approximately 90 ° relative to roof of skull, forming supraorbital crests; in this configuration there is no ventral groove for occipital artery. Supraorbital crests ornamented, composing a continuum with canthal crest; squamosal with a great mosaic of planes and processes, formed mainly by dorsal, vertical, and medial planes, medial plane blade-shaped, poorly developed, and projected to maxillary bone, horizontally oriented and poorly developed posteriorly; a large medial projection (otic plate) on top of squamosal; exoccipitals not fused one with another; a well developed prootic process that does not touch frontoparietal; a jagged or “scalloped” suture formed by articulation between pterygoid medial ramus (8) and the parasphenoid alae (7) (Fig. 6 B and 6 C; sensu Pramuk, 2006); neopalatine bone bearing a blade-like projection (Fig. 6 B and 6 C); dorsal surface of sphenethmoid covered completely by medial articulation of nasals and frontoparietals (6); nasals not expanded dorsolaterally (as they are in R. granulosa species group), and forming a transverse suture with frontoparietals; jaw articulation lies posterior to level of the fenestra ovalis; and with a complete temporal arcade (sensu Pramuk, 2006). Advertisement call: The recording was made at Centro de Instrução e Adestramento de Brasília, Brasília, Distrito Federal in August 15 2003, 22: 30 h. The air temperature was 17.1 ºC, with 73 % relative humidity. The specimen has SVL= 117.7 mm. The advertisement call graphics are shown in Figure 7. The calls were variable in duration (1.9– 12.4 s). Mean call rate was 9.8 ± 0.6 notes per second. (19–129 notes per call). Note rate and note interval were based on the means of 10 notes of the calls. The mean note rate was 44.7 (± 2.41) ms and the interval among note average 66.5 (± 2.17) ms; each note was formed by three pulses (Fig. 7 B). Dominant and fundamental frequencies were also determined from five notes from various parts of the calls. Mean dominant and fundamental frequencies were 839.0 (± 8.9) Hz and 421.0 (± 3.3) Hz, respectively. The advertisement call of R. cerradensis resembles that of other species in the R. marina group. Although the calls of the Rhinella marina species group are conservative, they present some differences. The notes per advertisement call of R. cerradensis is more variable than R. schneideri (33–40; Köhler et al., 1997), R. icterica (4–20; Heyer et al., 1990), R. poeppigii (10–45; De La Riva et al., 1996), R. arenarum (61–107; Straneck et al., 1993), R. marina (19–25; Crocroft et al., 2001), and R. rubescens (5–25) (present work). The number of pulses per note of the new species (three) is equal to those of R. schneideri (Köhler et al., 1997), and R. arenarum (Straneck et al., 1993), while in R. icterica they range from one to three (Heyer et al., 1990), three to five in R. poeppigii (De La Riva et al., 1996), five pulses in R. marina (Crocroft et al., 2001), and two pulses in R. rubescens (present work). The advertisement call of R. schneideri has a second power peak of 2100 Hz above the dominant frequency (Köhler et al., 1997). Rhinella icterica (Heyer et al., 1990), R. rubescens (Haddad et al., 1988), R. marina (Crocroft et al., 2001), R. poeppigii (De La Riva et al., 1996), R. arenarum (Straneck et al., 1993), and the new species lack the second power peak. However, the spectrogram of R. arenarum provided by Martin (1972) showed a second power peak above the dominant frequency. The re-analyzed call of R. icterica (Heyer et al., 1990) presented a lower fundamental and dominant frequencies of 324.9 (± 4.5) and 646.0 (± 0.2) Hz than R. cerradensis 421.0 (± 3.3) and 839.0 (± 8.9) Hz. Conversely, R. marina (Crocroft et al., 2001) showed higher fundamental and dominant frequencies of 559.9 and 1119.7 Hz (with no frequency variation among notes) among these three species (R. icterica, R. cerradensis, and R. marina). The advertisement call of R. arenarum (Straneck et al., 1993) presented a dominant frequency of 1348 (± 18.3) Hz, similar to R. rubescens, dominant fre

Universidade Federal de Uberlândia CHAPTER 2: The freshwater turtle Podocnemis expansa is found all over the Amazon basin and appears in almost all of its tributaries. The vertebrate skeleton is composed of cartilage and bone and represents the cell product of three different embryonic lineages. The cranium is formed by cells of the neural crest. Knowledge of the biological criterion for the sequence of bone formation is of highest importance for the understanding of ontogeny. Thus, the purpose of the present study was to establish normal stages of the formation and ossification sequence of the bony elements of the cranium in P. expansa, in the different stages of pre and post natal development. Embryos and nestlings were collected as from the 18th day of natural incubation. Morphometics occurred and the embryos were submitted to the diaphanous (clearing) technique and coloring of the bones. Neurocranium: in stage 19 the basisphenoid and basioccipital present the ossification center; in stage 20 the supraoccipital and opisthotic, in stage 21 the exoccipital and only in stage 24 the prootic. Dermatocranium: the squamosal, pterygoid and maxilla are the first elements to begin the ossification process, which occurs in stage 16. But most of these bone elements present ossification centers in stage 17, they are: frontal, jugal, postorbital, parietal, premaxilla, prefrontal, followed by the palatine and quadratojugal in stage 19 and last of all the vomer in stage 25. The esplancnocranium quadrate, in stage 23. Ossification of the mandible and hyoid apparatus: as much the dentary, coronoid and the surangular present ossification centers in stage 16 and the branchial body I in stage 17. The sequence and synchronization of the ossification in P. expansa show similarities, as well as differences, when compared with other chelonian species. It is evident that more quantitative studies are necessary to document the natural variability of formation of the bones. CHAPTER 3: The scleral ossicles are small bony plates located in the eyeball of many vertebrates and differ in their morphology, development and position within different groups of vertebrates. With the objective of investigating the development, number form and disposition of the scleral ossicles, in Podocnemis expansa 15 embryos were collected, as from the 18th day of natural incubation and 16 adult specimens, belonging to the LAPAS, from which the eyeballs were removed.The embryos and the eyeballs were submitted to the diafanization (Clearing) technique and coloration of the bones. On found out that the scleral ossicles, in Testudines species, have a square form, vary from 10 to 13 with 11,25 ± 0,93 and 11,43 ± 0,81, in the right eye and in the left eye, respectively, and they occupy a set position close to the front border of the sclera. The ossicles develop in an intramembranous form and begin to present ossification centers in stage 21, with ten ossification centers in the right eye and eight in the left eye. In stage 23, all ossicles of the sclerotic ring increase in length and approach the adjacent ossicles, forming a ring. In stage 26 the scleral ossicles have already presented themselves in the definitive disposition. CHAPTER 4: With the objective of investigating the formation sequence of the bony elements which form the spine, in the different stages of pre and post natal development in Podocnemis expansa, embryos and nestlings were collected, as from the 18th day of natural incubation. Morphometrics occurred and the embryos and nestlings were submitted to the diafanization and coloration technique of the bones and cartilages. The P. expansa spine comprises eight cervical vertebras, ten back vertebras, two sacral vertebras and twenty tale vertebras. In stage 15 the vertebras are still constituted by cartilage. In stage 17 the cartilage begins to be substituted by bone, and this takes place in the cranium-tail direction. In stage 19 all the back vertebras present ossification centers, as well as the two sacral vertebras. As from stage 20 the tail vertebras begin to present ossification centers. CHAPTER 5: Brazil has the richest fauna in all of Central and South America, but most of the information about reptiles is still incomplete. Podocnemis expansa, known popularly as the giant Amazon river turtle, is widely distributed through the basin of the Amazon and its tributaries. To investigate the sequence of the formation of the bone elements comprising the carapace in the various stages of pre- and postnatal development in P. expansa, embryos and newborns were collected, starting from the 18th day of natural incubation. Biometric measurements were taken and the embryos were subjected to the clearing technique and to bone and cartilage staining. The carapace is composed of 8 pairs of costal bones, 11pairs ribs, 7 neural bones, 11 pairs of peripheral bones, 1 nuchal bone that is part of the cranial portion of the carapace, 1 suprapigal bone, and 1 pigal bone that forms the caudal part of the carapace. In stage 16, ossification centers are present in the first seven pairs of ribs. In stage 17, the last pair of ribs begins to ossify. The bone expansions in the more cranial ribs begin in stage 19. The neural bones begin to ossify in stage 20, followed by the nuchal bone in stage 21. The first pair of peripheral bones presents ossification centers in stage 23, while the ossification process in the suprapigal and pigal scutes begins in stage 26. The bones that form the rigid armor of the carapace only begin to come together 46 days after hatching. CHAPTER 6: The Podocnemis expansa is known popularly as the giant Amazon river turtle and is considered one of the wild species more exploited most in zoo technology. With the objective of investigating the formation sequence of the bony elements which form the plastron/breastplate, in the different pre and post natal development stages, embryos and nestlings were collected, as from the 18th day of natural incubation. Morphometrics occurred and the embryos and nestlings were submitted to the diafanization and coloration technique of bones and cartilages. The epiplastron, endoplastron, hioplastron, hipoplastron, xifiplastron and the mesoplastron bones form the breastplate of these turtles. In stage 16 ossification centers are observed in the great majority of the breastplate bones. With regard to the retention of the alizarin the sequence is: first the hioplastron and the hipoplastron, then the endoplastron, followed by the xifiplastron and lastly the mesoplastron. The epiplastron bone showns itself with ossification center in stage 20 only. All these elements have independent ossification centers and they join together later. The closing of the breastplate only takes place seven months after hatching. CHAPTER 7: With the objective of investigating the sequence of bone formation of the breast bone and thoracic member, in the different stages of development be they pre or post natal Podocnemis expansa, embryos and newly born nestlings were collected, as from the 18th day of natural incubation. Morphometrics occurred and the embryos and nestlings were submitted to the diafanization and coloration technique of bones and cartilages. The arm skeleton (stylopodium) is composed by the humerus bone and the forearm skeleton (zeugopodium) consists of two bones, the ulna and the radius. The hand skeleton (autopodium) is formed by the following bones: carpus ulnare (UC), carpus central 2 (C2), carpus central 3 (C3), distal carpus bones (DCI, DCII, DCIII, DCIV and DCV), carpus pisiform (PC), intermediate carpus bone (IC), 5 metacarpus bones (MCI, MCII, MCIII, MCIV and MCV) and 14 phalanges, two in the first finger and three in each one of the other fingers, whose phalanx formula is 2:3:3:3:3. In stage 19, the scapula, the acromion and coracoid process already show ossification centers. In stage 16, ossification centers were observed in the humerus bone, followed by the ulna bone and, finally, the radius bone. In stage 23, the IC presents an ossification center, in stage 24, the whole distal row presented ossification centers, except for DC V. Still in that stage ossification centers were also verified in UC, CII and CIII. Only DCV showed ossification centers at the beginning of stage 25. PC is only seen 40 days after hatching. Retention of the stain in the metacarpi occurs in the following order: M III > M II = M IV > M I > M V. In stage 20, it is evident that, in all the metacarpi, the ossification centers progress in the direction of the epiphyses. The sequence of stain retention in the distal phalanges (DP) occurs in the order: DP III > DP II > DP IV > DP I > DP V. In the medial phalanges (MP), the stain retention sequence is: MP III > MP IV > MP II > MP V. Lastly, the proximal phalanges (PP) retain staining in the following sequence: PP I > PP III > PP IV > PP II > PP V. The differences and similarities in the synchronization of ossification of Podocnemis expansa and comparable species are evident. Thus, there is no osteogenetic pattern common to all chelonians, due to variations in the initial site of bone formation and the sequence of ossification. CHAPTER 8: With objective of investigating the sequence of bone formation of the pelvic girdle and pelvic members, in the different stages of pre and post-natal development in Podocnemis expansa, embryos and nestlings, as from the 18th day of natural incubation, were collected. Morphometrics took place and the embryos were submitted to the diafanization technique with coloration of bones and cartilages. The stylopodium area is composed by the femur bone, the zeugopodim consists of two bones, tibia and fibula and the autopodium is formed by the following bones: intermedium of tarso (IT), fibulare of tarso (FT), centrale of tarso (CT), distal bones of tarso (DTI, DTII, DT III and DTIV), 5 metatarsos (MtI, MtII, MtIII, MtIV and MtV) and 14 phalanges, two in the fifth finger and three in each one of the other fingers, whose phalange formula is 3:3:3:3:2. In stage 16 the femur, tibia and fibula present ossification centers in the diaphysis progressing from the epiphysis. In stage 19, the bones of the pelvic waist line show ossification centers. In the tarsal bones the DT I is the first to present ossification centers, at the beginning of stage 24, proceeding with IT, later CT and DT II at the end of stage 24, DT III and DT IV in stage 25. FT shows ossification center in the last embryonic stage, which is stage 26. Retention of the stain in the metatarsal occurs in the following order: M III > M II = M IV > M V > M I. In stage 20, it is evident that, in all the metatarsal, the ossification centers progress in the direction of the epiphyses. The sequence of stain retention in the distal phalanges (DP) occurs in the order: DP V > DP IV > DP III > DP II >DP I. In the medial phalanges (MP), the stain retention sequence is: MP III > MP IV > MP II > MP I. Lastly, the proximal phalanges (PP) retain staining in the following sequence: PP V > PP I > PP II > PP III > PP IV. The differences and similarities in the synchronization of ossification of Podocnemis expansa and comparable species are evident. Thus, there is no osteogenetic pattern common to all chelonians, due to variations in the initial site of bone formation and the sequence of ossification. CAPITULO 2: O cágado Podocnemis expansa está distribuído pela bacia amazônica e ocorre em quase todos os seus afluentes. O esqueleto dos vertebrados é composto de cartilagem e osso e representa o produto de células de três linhagens embrionárias distintas. O crânio é formado por células da crista neural. O conhecimento do critério biológico para a seqüência de formação óssea é de suma importância para o entendimento da ontogenia. Assim, o propósito do presente estudo foi estabelecer etapas normais da formação da seqüência de ossificação dos elementos ósseos do crânio em P. expansa, nos diferentes estágios de desenvolvimento pré e pós-natal. Coletaram-se embriões a partir do 18º dia de incubação natural. Efetuou-se morfometria e os embriões foram submetidos à técnica de diafanização e coloração dos ossos. Neurocrânio: no estágio 19 o basisfenóide e o basioccipital apresentam centro de ossificação; no estágio 20 o supra-occipital e o opistótico, no estágio 21 o exoccipital e somente no estágio 24 o proótico. Dermatocrânio: o esquamosal, o pterigóide e a maxila são os primeiros elementos a iniciar o processo de ossificação, isso ocorre no estágio 16. Mas a maioria desses elementos ósseos apresenta centros de ossificação no estágio 17, são eles: frontal, jugal, pós-orbital, parietal, pré-maxila, pré-frontal, seguido do palatino e quadradojugal no estágio 19 e por último o vômer no estágio 25. O osso quadrado do esplancnocrânio, no estágio 23. Ossificação da mandíbula e aparelho hióide: tanto o dentário, coronóide e o supra-angular apresentam centros de ossificação no estágio 16 e o corpo branquial I no estágio 17. A seqüência e a sincronização da ossificação em P. expansa exibem similaridades, bem como diferenças, quando comparada com outras espécies de Testudines. Fica evidente que mais estudos quantitativos são necessários para documentar a variabilidade natural de formação dos ossos. CAPITULO 3: Os ossículos da esclera são pequenas placas ósseas localizadas no globo ocular de muitos vertebrados e diferem em sua morfologia, desenvolvimento e posição dentro de diferentes grupos de vertebrados. Com objetivo de investigar o desenvolvimento, o número, a forma e a disposição dos ossículos da esclera, em Podocnemis expansa coletou-se 15 embriões, a partir do 18º dia de incubação natural e 16 espécimes adultos, pertencentes ao acervo do LAPAS, dos quais foram removidos os globos oculares. Os embriões e os globos oculares foram submetidos à técnica de diafanização e coloração dos ossos. Apurou-se que os ossículos da esclera, nessa espécie de Testudines, possuem forma quadrangular, variam de 10 a 13 com 11,25 ± 0,93 e 11,43 ± 0,81, no olho direito e no olho esquerdo, respectivamente, e ocupam posição fixa, próximos à borda anterior da esclera. Os ossículos se desenvolvem de forma intramembranosa e começam a apresentar centros de ossificação no estágio 21, com dez centros de ossificação no olho direito e oito no olho esquerdo. No estágio 23, todos os ossículos do anel esclerótico aumentam em comprimento e se aproximam dos ossículos adjacentes, formando um anel. No estágio 26 os ossículos da esclera já se apresentam na disposição definitiva. CAPITULO 4: Com objetivo de se investigar a seqüência de formação dos elementos ósseos que formam a coluna vertebral, nos diferentes estágios de desenvolvimento pré e pós-natal em Podocnemis expansa, coletaram-se embriões e filhotes, a partir do 18º dia de incubação natural. Efetuou-se morfometria e os embriões e filhotes foram submetidos à técnica de diafanização e coloração dos ossos e cartilagens. A coluna vertebral de P. expansa compreende oito vértebras cervicais, dez vértebras costais, duas vértebras sacrais e vinte vértebras caudais. No estágio 15 as vértebras ainda são constituídas por cartilagem. No estágio 17 a cartilagem começa a ser substituída por osso, e ocorre no sentido crânio-caudal. No estágio 19 todas as vértebras costais apresentam centros de ossificação, além das duas vértebras sacrais. A partir do estágio 20 as vértebras caudais começam a apresentar centros de ossificação. CAPITULO 5: O Brasil tem a fauna mais rica de toda a América Central e do Sul, mas a maioria das informações sobre répteis é ainda preliminar. A Podocnemis expansa, conhecida popularmente como tartaruga-da-amazônia, é largamente distribuída pela bacia amazônica e seus afluentes. Com objetivo de investigar a seqüência de formação dos elementos ósseos que formam a carapaça, nos diferentes estágios de desenvolvimento pré e pós-natal, em P. expansa coletaram-se embriões e filhotes recém-nascidos, a partir do 18º dia de incubação natural. Efetuou-se morfometria e os embriões foram submetidos à técnica de diafanização e coloração dos ossos e cartilagens. A carapaça é composta por 8 pares de ossos costais, 11 pares de costelas, 7 ossos neurais, onze pares de ossos periféricos, 1 osso nucal que forma a parte cranial da carapaça, 1 osso supra-pigal e 1 osso pigal que forma a parte caudal da carapaça. No estagio 16 estão presentes centros de ossificação do segundo ao sétimo par de costelas. No estágio 19, iniciam-se expansões ósseas nas costelas mais craniais. Os ossos neurais começam a se ossificar no estágio 20, seguindo do osso nucal no estagio 21. No estágio 23 o primeiro par de ossos periféricos apresentam centros de ossificação. No estágio 26, os ossos supra-pigal e o pigal iniciam o processo de ossificação. A aproximação dos ossos que formam a armadura rígida da carapaça, só ocorre depois de 46 dias da eclosão. CAPITULO 6: A Podocnemis expansa é conhecida popularmente como tartaruga-da- Amazônia e é considerada uma das espécies selvagens mais exploradas zootecnicamente. Com objetivo de se investigar a seqüência de formação dos elementos ósseos que formam o plastrão, nos diferentes estágios de desenvolvimento pré e pós-natal, coletaram-se embriões e filhotes, a partir do 18º dia de incubação natural. Efetuou-se biometria e os embriões e filhotes foram submetidos à técnica de diafanização e coloração dos ossos e cartilagens. Os ossos epiplastrão, endoplastrão, hioplastrão, hipoplastrão, xifiplastrão e o mesoplastrão formam o plastrão desse testudines. No estágio 16 observam-se centros de ossificação na grande maioria dos ossos do plastrão. De acordo com a retenção do corante alizarina a seqüência é: primeiro o hioplastrão e o hipoplastrão, depois o endoplastrão, seguido do xifiplastrão e por último o mesoplastrão. O osso epiplastrão mostra-se com centro de ossificação apenas no estágio 20. Todos esses elementos possuem centros de ossificação independentes e se unem posteriormente. O fechamento do plastrão só ocorre depois de sete meses após a eclosão. CAPITULO 7: Com objetivo de investigar a seqüência de formação dos ossos da cintura peitoral e do membro torácico, nos diferentes estágios de desenvolvimento pré e pós-natal em Podocnemis expansa, coletou-se embriões e filhotes recémnascidos, a partir do 18º dia de incubação natural. Efetuou-se biometria e os embriões e filhotes foram submetidos à técnica de diafanização e coloração dos ossos e cartilagens. O esqueleto do braço (estilopódio) é composto pelo osso úmero e o esqueleto do antebraço (zeugopódio) compreende dois ossos, a ulna e o rádio. O esqueleto da mão (autopódio) é formado pelos ossos: ulnar do carpo (UC), central do carpo 2 (C2), central do carpo 3 (C3), ossos distais do carpo (DCI, DCII, DCIII, DCIV e DCV), pisiforme do carpo (PC), osso intermédio do carpo (IC), 5 ossos metacarpos (MCI, MCII, MCIII, MCIV e MCV) e 14 falanges, duas no primeiro dedo e três em cada um dos demais dedos, cuja fórmula falângica é 2:3:3:3:3. No estágio 19, a escápula, o processo acromial e o coracóide já mostram centros de ossificação. No estágio 16, observaram-se centros de ossificação no osso úmero, seguido pelo osso ulna e, finalmente, o osso rádio. No estágio 23, o IC apresenta centro de ossificação, no estágio 24, toda a fileira distal do carpo apresentou centros de ossificação, exceto do DC V. Ainda nesse estágio também se verificaram centros de ossificação nos UC, CII e CIII. O DCV só mostrou centros de ossificação no início do estágio 25. O PC somente é visto 40 dias após a eclosão. A retenção do corante nos metacarpos ocorre na seguinte ordem: MCIII > MCII = MCIV > MCI > MCV. No estágio 20 torna-se evidente que, em todos os metacarpos, os centros de ossificação progridem em direção às epífises. A seqüência de retenção de corante nas falanges distais (FD) ocorre na ordem: FD III > FD II > FD IV > FD I > FD V. Nas falanges médias (FM): FM III > FM IV > FM II > FM V. E, finalmente, nas falanges proximais (FP): FP I > FP III > FP IV > FP II > FP V. Diferenças e semelhanças na sincronização de ossificação são evidentes entre Podocnemis expansa e as espécies comparadas. Assim, não existe um padrão de osteogênese para todos os Testudines, devido à variação do local inicial de formação óssea e da seqüência de ossificação. CAPITULO 8: Com objetivo de investigar a seqüência de formação óssea da cintura pelvina e dos membros pelvinos, nos diferentes estágios de desenvolvimento pré e pós-natal em Podocnemis expansa, coletou-se embriões e filhotes, a partir do 18º dia de incubação natural. Efetuou-se biometria e os embriões foram submetidos à técnica de diafanização com coloração dos ossos e cartilagens. A região do estilopódio é composta pelo osso fêmur, o zeugopódio compreende dois ossos, tíbia e fíbula e o autopódio é formado pelos ossos: intermédio do tarso (IT), fibular do tarso (FT), central do tarso (CT), distais do tarso (DTI, DTII, DT III e DTIV), 5 metatarsos (MtI, MtII, MtIII, MtIV e MtV) e 14 falanges, duas no quinto dedo e três em cada um dos demais dedos, cuja fórmula falângica é 3:3:3:3:2. No estágio 16 o fêmur, tíbia e fíbula apresentam centros de ossificação na diáfise progredindo para as epífises. No estágio 19, os ossos da cintura pelvina mostram centros de ossificação. Nos ossos do tarso o DT I é o primeiro a apresentar centros de ossificação, no início do estágio 24, seguindo com IT, depois o CT e o DT II no final do estágio 24, DT III e DT IV no estágio 25. O FT mostra centro de ossificação no último estágio embrionário, estágio 26. A retenção do corante nos metatarsos ocorre na seguinte ordem: MtIII>MtII=MtIV>MtV>MtI. No estágio 20 torna-se evidente que, em todos os metatarsos, os centros de ossificação progridem em direção às epífises. A seqüência de retenção de corante nas falanges distais (FD) ocorre na ordem: FD V > FD IV > FD III > FD II >FD I. Nas falanges médias (FM): FM III > FM IV > FM II > FM I. Nas falanges proximais (FP): FP V > FP I > FP II > FP III > FP IV. As diferenças e semelhanças na sincronização de ossificação são evidentes entre Podocnemis expansa e as espécies comparadas. Assim, não existe um padrão de osteogênese para os Testudines, devido à variação do local inicial de formação óssea e da seqüência de ossificação. Mestre em Ciências Veterinárias

Epipedobates flavopictus is a diurnal, aposematic dendrobatid with a wide distribution in seasonal wet tropical regions of Brazil. We describe the daily period of vocalization, advertisement call, courtship behavior, and tadpole of E. flavopictus from a previously unknown population in southern Goiás and compare theses features with that of other populations. Studies were carried out in November (2004) and February (2005). We counted the number of calling males and duration of calling bouts in the morning and evening periods. The advertisement call was recorded with a digital recorder. Tadpole description was based on specimens collected in pools. Males called from well-illuminated sites such as rocky fields, rain channels, and borders of riverine forests. In November, males vocalized daily during two distinct periods, between 0430 and 1000 h and between 1630 and 2000 h. Morning temperature varied between 20-23ºC and humidity from 79-89%; during evening varied between 24-27ºC and 54-82%. In the middle of the day, temperature reached 36ºC and humidity 40%. During the morning, call activity was almost uninterrupted, in the evening calling bouts lasted around 9 min. In February, even with the occurrence of rainfall and temperatures similar to that of November no frog vocalized. The advertisement call is composed by a single note with 7 8 pulses with frequency ascending slightly from 3.20 to 4.05 kHz. Note duration was 144 ms, and between note intervals is 292 ms. Notes are given at a rate of 139 per minute. Upon observing the female, the male began to emit courtship call. The female approached the male and touched him laterally with her snout. The male moved forward and raised his hindquarters by stretching his hind legs. While leading the female, the male continued to give courtship and advertisement calls. The male clasped the female in axillary amplexus, and the pair entered a hole in a bank. The female deposited eggs on the surface of the soil, spreading them in groups. The egg clutches had eggs in at least two developmental stages, recently deposited and with embryos. Tadpoles were found in small, shallow rocky pools along a permanent stream, in well-illuminated sites at the forest border. The tadpole had the dorsal fin arched, not extending onto body. E. flavopictus appears to be unique among dendrobatids by using open areas subject to high temperatures and low humidity. Reproductive activity of the species ends before the end of the rainy season, possibly to avoid loss of tadpoles during months with unpredictable rainfall. The studied call had more notes and shorter between-call intervals than those described from other populations. The free-living tadpoles we describe differ from those of other populations by having the dorsal fin reduced. Epipedobates flavopictus é um dendrobatídeo diurno, aposemático que tem ampla distribuição em regiões tropicais do Brasil. Descrevemos o período diário de atividade de canto, canto de anúncio, comportamento de corte e girino de E. flavopictus de uma população previamente desconhecida do sul de Goiás e comparamos essas características às de outras populações. O estudo foi realizado em novembro de 2004 e fevereiro de 2005. Comparamos o número de machos em atividade de canto e a duração dos períodos de canto da manhã e da tarde. O canto foi gravado com aparelho digital. A descrição dos girinos foi feita com base em espécimes coletados em poças. Os machos vocalizavam em sítios bem iluminados tais como ambientes rupestres e borda de matas ribeirinhas. Em novembro, os machos vocalizavam diariamente em dois períodos diferentes do dia; entre 0430 e 1000 h e entre 1630 e 2000 h. Durante a manhã, a temperatura variou entre 20 e 23 ºC e a umidade entre 79 e 89%; durante a tarde variou entre 24 e 27 ºC e 54 e 82%. No meio do dia, a temperatura chegou a 36 ºC e a umidade a 40%. Durante a manhã, a atividade de canto era quase ininterrupta, e a tarde os períodos de canto duravam cerca de 9 min. Em fevereiro, apesar da ocorrência de chuvas e temperatura/umidade similares as de novembro, não houve atividade de canto. O canto de anúncio é composto por uma única nota com 7-8 pulsos, com freqüência ascendente de 3,20 a 4,05 kHz. A duração da nota é de 144 ms com intervalos de 292 ms. As notas são emitidas a uma taxa de 139 notas por minuto. Uma vez visualizada a fêmea, o macho começa a emitir canto de corte. A fêmea se aproxima do macho e o toca na lateral do corpo com o focinho. O macho se move pra frente e levanta o seu quarto traseiro esticando suas pernas. Enquanto conduzindo a fêmea, o macho continuava a emitir cantos de corte e de anuncio. O macho amplexou a fêmea axilarmente, e o par entrou num buraco no barranco. A fêmea depositou os ovos na superfície do solo, espalhando-os em grupos. Uma desova examinada tinha ovos em dois estágios diferentes de desenvolvimento: recém depositados e com embriões. Os girinos foram encontrados em pequenas poças rasas em rochas ao longo de um riacho permanente, em sítios bem iluminados na borda da floresta. Os girinos tinham a nadadeira dorsal arqueada, a qual não se estendia para o corpo. Aparentemente, E. flavopictus é o único entre os dendrobatídeos por usar áreas abertas sujeitas a altas temperaturas e baixas umidades. A atividade reprodutiva da espécie termina antes do fim da estação chuvosa, possivelmente para evitar perda de girinos durante meses com chuvas imprevisíveis. O canto estudado tinha mais notas e intervalos de cantos mais curtos que aqueles descritos de outras populações. O girino de vida livre que descrevemos difere do de outras populações ter a nadadeira dorsal reduzida. Mestre em Ecologia e Conservação de Recursos Naturais