A New Species of the Bokermannohyla circumdata Group (Anura: Hylidae)

from the Coastal Forests of Bahia, Northeastern Brazil

Marcelo F. Napoli1,2 and Bruno V. S. Pimenta3

We describe a new species of tree frog related to the Bokermannohyla circumdata group, the northernmost record fromAtlantic Forest remnants in eastern Bahia, northeastern Brazil, within regions known as Vale do Jiquirica, Baixo Sul, andLitoral Sul. The new species is diagnosed by medium size (snout–vent length 46.8–64.1 mm), head longer than wide,small tympanum, posterior surfaces of thighs with dark brown transverse stripes without additional thin stripesbetween them, vocal slits in adult males, and distinct advertisement call with a single note composed of 30–110 pulsesdistributed within 2–9 pulse groups, similar to the bleating of a goat. The new species is morphologically similar to B.hylax and B. lucianae, but is readily distinguished by the presence of barred posterior surfaces on the thighs without thinstripes between them (immaculate in B. lucianae; thin stripes intercalated with wider stripes in B. hylax) and by itsdistinct vocalization.

Nos descrevemos uma nova especie de perereca relacionada ao grupo de Bokermannohyla circumdata, o registro maissetentrional para remanescentes de Floresta Tropical Atlantica do leste da Bahia, nordeste do Brasil, em regioesconhecidas como Vale do Jiquirica, Baixo Sul e Litoral Sul. A nova especie e diagnosticada principalmente pelo tamanhomedio (comprimento rostro-cloacal 46,8–64,1 mm), cabeca mais longa que larga, tımpano pequeno, superfıciesposteriores da coxa com faixas transversais castanho escuras sem faixas finas adicionais entre elas, fendas vocais emmachos adultos e canto de anuncio distinto, com uma unica nota composta por 30–110 pulsos distribuıdos em 2–9grupos de pulsos, o qual soa como um balido de bode. A nova especie e morfologicamente similar a B. hylax e B. lucianae,mas e prontamente distinguida por apresentar superfıcies posteriores das coxas barradas sem faixas finas entre elas(imaculadas em B. lucianae; faixas finas intercaladas com faixas mais largas em B. hylax) e por sua vocalizacao distinta.

THE Bokermannohyla circumdata species group current-ly comprises 17 species that occur in mountainstream habitats within the Tropical Atlantic Forest of

Brazil (B. ahenea, B. astartea, B. caramaschii, B. carvalhoi, B.circumdata, B. diamantina, B. feioi, B. gouveai, B. hylax, B.ibitipoca, B. izecksohni, B. lucianae, B. luctuosa, B. nanuzae, B.ravida, B. sazimai, and B. vulcaniae) and is diagnosed by thepresence of the following combination of traits: browndorsum (in live and preserved specimens), simple (notdichotomized) transverse bands on the posterior surface ofthighs and flanks, single (not bifid) large and sharpprepollex, and hypertrophied forearm in adult males(Napoli and Junca, 2006). Faivovich et al. (2005) hadpreviously diagnosed this group by 52 transformations innucleotide sequences (ribosomal genes and protein codinggenes from the nucleus and mitochondrion) and by thepresence of (usually thin) dark vertical stripes on theposterior surface of thigh, the latter a putative morpholog-ical synapomorphy. During surveys of anurans in coastalAtlantic Forest remnants in eastern Bahia, northeasternBrazil, within regions known as Vale do Jiquirica, Baixo Sul,and Litoral Sul, we found a new species that we describeherein and assign to the B. circumdata group.

MATERIALS AND METHODS

We examined 15 morphometric characters following Napoli(2005): SVL (snout–vent length), HL (head length), HW(head width), ED (eye diameter), UEW (upper eyelid width),IOD (interorbital distance), IND (internarial distance), END

(eye–nostril distance), NSD (nostril to tip of snout distance),TD (tympanum diameter), THL (thigh length), TL (tibialength), FL (foot length), 3FD (third finger disk diameter),and 4TD (fourth toe disk diameter). Measurements weretaken with the help of an ocular micrometer in an Olympusstereomicroscope, except for SVL, HL, HW, THL, TL, and FL,which were measured with a vernier caliper to the nearest0.1 mm. Drawings were made using a Leica MZ6 stereomi-croscope with a drawing tube. Webbing formula notationfollowed Savage and Heyer (1967), as modified by Myers andDuellman (1982). Standards for dorsal outline of snout andsnout profile followed Heyer et al. (1990). Institutionalabbreviations are as listed at http://www.asih.org/codons.pdf, with the addition of MZUEFS (Museu de Zoologia daUniversidade Estadual de Feira de Santana, Bahia, Brazil) andUFBA (Museu de Zoologia da Universidade Federal da Bahia,Salvador, Bahia, Brazil).

Advertisement calls of the new species were recorded fromthe following Brazilian municipalities in the State of Bahia:Amargosa, Serra do Timbo, 13u049S, 39u389W, 18–19 July2007, M. F. Napoli, P. M. Fonseca, and R. O. Abreu, using aMarantz Portable Cassette Recorder PMD222 with a Senn-heiser ME66 directional microphone; Camamu, Projeto deAssentamento Zumbi dos Palmares, 14u019S, 39u099W, 11–12 December 2000, Bruno V. S. Pimenta and Paulo H. C.Cordeiro, using a Sony DAT TCD-D8 tape recorder with aSennheiser ME66/K6 directional microphone set; Valenca,RPPN Agua Branca, 13u229S, 39u049W, 6–9 September 2004,Bruno V. S. Pimenta, using a Sony DAT TCD-D8 taperecorder with a Sennheiser ME66/K6 directional micro-

1 Museu de Zoologia, Departamento de Zoologia, Instituto de Biologia, Universidade Federal da Bahia, Campus Universitario, Rua Barao deJeremoabo 147, Ondina, 40170-115 Salvador, Bahia, Brazil; E-mail: napoli@ufba.br. Send reprint requests to this address.

2 Associate Researcher, Departamento de Vertebrados, Museu Nacional/Universidade Federal do Rio de Janeiro, Quinta da Boa Vista, SaoCristovao, 20940-040 Rio de Janeiro, Rio de Janeiro, Brazil.

3 BBM Consultoria Ambiental Ltda., Av. Prudente de Morais 621/807, Cidade Jardim, 30380-000, Belo Horizonte, Minas Gerais, Brazil; E-mail:bvergueiropimenta@gmail.com.

Submitted: 29 November 2008. Accepted: 11 May 2009. Associate Editor: D. Kizirian.F 2009 by the American Society of Ichthyologists and Herpetologists DOI: 10.1643/CH-08-224

Copeia 2009, No. 4, 674–683

phone set; Wenceslau Guimaraes, Estacao Ecologica Esta-dual Nova Esperanca, Mata do Jorge and Mata do Pequi,13u369S, 39u439W, 29 August 2000, Paulo H. C. Cordeiro,using a Sony DAT TCD-D8 tape recorder with a SennheiserME66/K6 directional microphone set. For detailed acousticcomparisons, we used advertisement calls of Bokermannohylahylax recorded from its type locality (Estacao Biologica deBoraceia, State of Sao Paulo, 23u389S, 45u529W), availablefrom the audio supplement of Heyer et al. (1990), and fromthe Municipality of Guaramirim, State of Santa Catarina,Brazil (26u289S, 49u009W); for B. diamantina, B. feioi, B.ibitipoca, B. lucianae, and B. luctuosa we analyzed the sametapes used in their original descriptions (Pombal andHaddad, 1993; Napoli and Pimenta, 2003; Napoli andCaramaschi, 2004; Napoli and Junca, 2006); for theremaining species of the B. circumdata group for which callswere available (B. nanuzae and B. sazimai) we examined onlypublished data (Bokermann and Sazima, 1973; Cardoso andAndrade, 1982). Calls were digitized at a sampling rate of8 kHz. Temporal parameters, number of pulses, number ofnotes, and frequency amplitude were measured or countedon waveforms with Avisoft SAS-Lab Light for Windows(version 3.74). Dominant frequencies were obtained withSound Ruler Acoustic Analysis, version 0.9.6.0. Audiospec-trograms were made with Fast Fourier Transform length(FFT) of 256, overlap 87.5%, frame 100%, window Flat Top,and frequency resolution of 161 Hz. Nine call parameterswere measured: note number, note duration (s), inter-noteduration (s), note rate (notes/s), pulse number, number ofpulse groups, note pulse rate (pulses/s), call dominantfrequency (kHz), and frequency amplitude. Note pulse ratewas calculated for each note using the equation: pulse rate 5

(pulse number – 1)/note duration (s); note rate with theequation: note rate 5 note number/call duration (s). Theterminology used for the description of advertisement callsfollowed Duellman and Trueb (1994).

Bokermannohyla capra, new speciesFigures 1–3; Tables 1, 2

Hyla hylax.—Napoli, 2000:110 (in part, occurrence inUrucuca, Bahia State, Brazil).

Bokermannohyla hylax.—Junca, 2006:4–6, 11, 17, table 3, fig.3d (occurrence in Serra da Jiboia, Bahia State, Brazil).

Holotype.—UFBA 7492, adult male, Brazil, State of Bahia,Municipality of Amargosa, Serra do Timbo, 13u049S,39u389W, 800 m elevation, 18–19 July 2007, M. F. Napoli,P. M. Fonseca, and R. O. Abreu.

Paratypes.—Brazil: Bahia: UFBA 6415, adult female, UFBA6425, 6437, 6444, 6465–66, adult males, Municipality ofAmargosa, Serra do Timbo, 13u049S, 39u389W, 800 m, 13–22January 2007, P. M. Fonseca, T. F. Miranda, and B. Hamdan;UFBA 7483–88, 7490–91, adult males, UFBA 7489, adultfemale, collected with holotype; MNRJ 29183–84, adultfemales, Municipality of Camamu, Projeto de AssentamentoZumbi dos Palmares, 14u019S, 39u099W, 120 m, 11–12December 2000, B. V. S. Pimenta and P. H. C. Cordeiro;MZUEFS 653–55, Municipality of Elısio Medrado, 12u569S,39u319W, 800 m, 17 June 2001, F. A. Junca; UFBA 7000,Municipality of Itacare, 14u169S, 38u599W, 8 September2006; MZUEFS 2879–80, Municipality of Itubera, ReservaMichelin, 13u509S, 39u109W, 21 September 2007, F. A. Junca,F. Camurugi, and I. S. Nunes; MNRJ 29178–79, Municipality

of Nilo Pecanha, Fazenda Sao Joao, 13u429S, 39u149W,200 m, 5 September 2000, B. V. S. Pimenta and R. V. Lopes;MZUEFS 1268–70, Serra da Jiboia, 12u529S, 39u289W, 800 m,16 December 2001, F. A. Junca; MNRJ 25587–89, Munici-pality of Urucuca, Fazenda Caititu, 14u259S, 39u049W,140 m, 19 July 2000, B. V. S. Pimenta and R. V. Lopes;MNRJ 36940, adult female, MNRJ 36941–45, adult males,Municipality of Valenca, RPPN Agua Branca, 13u229S,39u049W, 6–9 September 2004, B. V. S. Pimenta and R. V.Lopes; MNRJ 29175–77, 29180–81, adult males, MNRJ29182, juvenile, Municipality of Wenceslau Guimaraes,Estacao Ecologica Estadual Nova Esperanca, Mata do Jorgeand Mata do Pequi, 13u369S, 39u439W, 470 m, 29 August2000, B. V. S. Pimenta and R. V. Lopes.

Diagnosis.—Diagnosed from all congeners by the combina-tion of medium to large size (SVL 46.8–64.1 mm in adultspecimens); head longer than wide, its length 2.5–3.0 timessmaller than SVL; tympanum small (TD/SVL 0.05–0.06);dorsum medium brown to pale cream, without distincttransverse dark brown bars; flanks pale cream, with few, orwithout, thin brown perpendicular stripes; anterior, superior,and posterior surfaces of thighs with dark brown transversethin stripes, not dichotomized; transverse brown bars onflanks and thighs without additional thinner stripes betweenthem; vocal slits present in adult males; crista ventralis ofhumerus not well developed, not producing an extensivehook-like humeral crest; single (not bifid) large and sharpprepollex; advertisement call consisting of a single notecomprised by 30–110 pulses distributed within 2–9 pulsegroups, which sounds like a bleating of a goat (Capra aegagrus).

Bokermannohyla capra is larger (male SVL 46.8–56.8 mm;female SVL 58.9–64.1 mm) than B. astartea, B. feioi, B.ibitipoca, B. nanuzae, B. oxente, and B. sazimai (combinedspecies SVL: males 30.2–43.4 mm; females 39.5–44.5 mm);males of B. capra are larger than males of B. claresignata andB. clepsydra (combined species SVL 36.0–42.0 mm), smallerthan males of B. caramaschii, B. carvalhoi, B. circumdata, B.gouveai, B. luctuosa, and B. martinsi (combined species SVL57.5–71.7 mm), and much smaller than males of B.alvarengai (SVL 60.7–140.9 mm); females of B. capra arelarger than females of B. ravida, B. vulcaniae, and B. itapoty(combined species SVL 38.0–53.3 mm), and much smallerthan females of B. alvarengai (SVL 79.9–87.6 mm). Tympa-num diameter of B. capra (males and females TD/SVL 0.05–0.06) is smaller than in B. vulcaniae (TD/SVL: males 0.09–0.10; females 0.09–0.11), B. caramaschii, B. luctuosa (com-bined species TD/SVL: males 0.08–0.10; females 0.07–0.09),and B. circumdata (TD/SVL: males 0.07–0.10; females 0.07–0.09). Head longer than wide in B. capra, wider than long inB. alvarengai, B. caramaschii, B. circumdata, B. claresignata, B.gouveai, B. ibitiguara, B. itapoty, B. langei, B. luctuosa, B.pseudopseudis, and B. vulcaniae. Snout rounded in dorsaloutline in B. capra, truncate in B. alvarengai, B. ibitiguara, andB. itapoty. Presence of vocal slits in adult males distinguishesB. capra from B. caramaschii and B. izecksohni, which lacksuch structures. Dorsum brown to pale cream, uniform orwith weak transverse darker brown stripes in B. capra; creamwith reticulated thin brown stripes and random brown dotsover it in B. sazimai; immaculate with dorsal surfaces goldenbrown in B. ahenea and B. astartea; blackish brown to lightbrown with distinct dark transverse brown stripes in B.caramaschii, B. circumdata, B. feioi, B. hylax, B. luctuosa, andB. ravida; brown intensively maculated with dark brown

Napoli and Pimenta—New Bokermannohyla from Bahia, Brazil 675

blotches somewhat anastomosed in B. oxente, B. itapoty, B.pseudopseudis, and B. saxicola. Flanks usually immaculate orscarcely barred with thin stripes (not dichotomized) in B.capra, distinctly barred in B. caramaschii, B. circumdata, B.feioi, B. hylax, B. luctuosa, B. ravida, and B. vulcaniae; stripesintercalated with wider transverse brown bars in B. hylax (B.capra lacks such pattern); stripes reticulated or dichotomizedin B. ibitiguara, B. pseudopseudis, and B. saxicola; flanks withbrown spots in B. clepsydra and B. claresignata. Absence of adorsolateral blackish brown stripe from tip of snout toinguinal region distinguishes B. capra from B. feioi, whichpossesses such stripe. The presence of a not well-developedcrista ventralis in humerus and a single large and sharpprepollex in males distinguish B. capra from B. langei and B.martinsi, in which the crista ventralis produces an extensivehook-like humeral crest and the prepollex is bifid (Napoli,2000; Caramaschi et al., 2001), and from B. claresignata andB. clepsydra, in which prepollex is poorly developed.Anterior, superior, and posterior surfaces of thighs with 8–11 dark thin transverse brown stripes in B. capra; anteriorand posterior surfaces immaculate in B. alvarengai, B.claresignata, B. clepsydra, B. feioi, B. nanuzae, B. lucianae,and B. oxente, with thin and fragmented stripes in B.carvalhoi, with thin brown stripes intercalated with widertransverse brown bars in B. hylax, with dark anastomosedvertical stripes (irregular, X-shaped and Y-shaped) on dorsaland posterior surfaces in B. vulcaniae; dorsal surfaces ofthighs with 4–6 large dark brown bars in B. alvarengai, B.ibitiguara, B. oxente, B. pseudopseudis, and B. saxicola, andwith brown spots in B. clepsydra and B. claresignata.Bokermannohyla capra is only found inside Atlantic forestremnants, whereas B. alvarengai, B. ibitiguara, B. itapoty, B.langei, B. martinsi, B. oxente, B. pseudopseudis, and B. saxicola

occur in rocky mountain fields and/or along riparian forests.Advertisement call distinguishes B. capra from all otherspecies of Bokermannohyla for which such data is available.

Description of holotype.—Head longer than wide, its widthand length 2.7 and 2.9 times smaller than snout–ventlength, respectively. Internarial distance smaller than eye–nostril distance and eye diameter. Eye diameter larger thaneye–nostril distance. Snout rounded in dorsal outline and inprofile. Canthus rostralis rounded. Loreal region slightlyconcave. Tympanum small, nearly circular, its width 1.7times smaller than eye diameter. Supratympanic fold distinct,covering upper border of the tympanum. Nostril superolat-eral. Internarial region slightly furrowed. Vomerine teethpresent in two angular patches between choanae. Tonguecordiform. Vocal sac single, subgular. Vocal slits large.Forearm hypertrophied, with weak dermal fringe. Thirdfinger disc diameter larger than fourth toe disk. Palmarsubarticular tubercles rounded. Distal tubercle of fourthfinger bifid. Palmar supernumerary tubercles present. Palmartubercle divided. Prepollex large, curved. Nuptial asperitiesnot present. Modal webbing formulae notation, I 2L–2L II2–3 III 2K–2 IV. Thigh and tibia lengths equal, their sumlarger than snout–vent length. Calcar distinct, small. Plantarsubarticular tubercles rounded. Plantar supernumerary tu-bercles distinct. Inner metatarsal tubercle ovoid. Modalwebbing formulae notation, I 1+–22 II 1–2+ III 1+–2K IV 2+–1+ V. Throat, chest, and dorsal and ventral surfaces of armssmooth. Belly and ventral surfaces of thighs glandular.

Measurements of holotype.—Snout–vent length 50.3 mm,head length 18.1 mm, head width 17.2 mm, eye diameter5.2 mm, eye–nostril distance 4.8 mm, tympanum diameter

Fig. 1. Holotype of Bokermannohyla capra, UFBA 7492 (SVL 50.3 mm), adult male, in dorsal (left) and ventral (right) views.

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2.9 mm, upper eyelid width 4.2 mm, interorbital distance5.8 mm, internarial distance 3.2 mm, nostril to tip of snoutdistance 2.9 mm, thigh length 25.7 mm, tibia length25.7 mm, foot length 35.0 mm, third finger disk diameter2.6 mm, fourth toe disk diameter 2.3 mm.

Color of holotype in preservative.—Dorsal ground color lightto tan brown, with scattered dark brown blotches. Two weaktransverse brown bars on urostyle region. Flanks pale cream

with three almost indistinct thin perpendicular brownstripes. Forearms with one weak dark brown transversestripe. Brachium with two thin brown transverse stripes overcream background color. Anterior, superior, and posteriorsurfaces of thighs with 9–10 thin dark brown transversestripes of same width and appearance, over pale creambackground color. Shanks, tarsus, and feet with up to fiveindistinct transverse brown stripes. Dark brown vertebralstripe present from tip of snout almost reaching sacrum.

Fig. 2. Holotype of Bokermannohyla capra, UFBA 7492, adult male. Dorsal (A) and lateral (B) views of head; ventral views of hand (C) and foot (D).Upper and lower horizontal lines equal 2.0 mm.

Napoli and Pimenta—New Bokermannohyla from Bahia, Brazil 677

Loreum and canthus rostralis of same dorsal backgroundcolor, immaculate. White lichenose marks over cloacalopening and heels. Ventral surfaces pale cream.

Color of holotype in life.—At night, dorsum reddish brownwith weakly visible dark transverse bands, few scatteredcream pinpoint spots, and dark brown blotches. Dark brownmid-dorsal stripe from tip of snout to near cloacal opening,where it reaches its smallest width. Flanks pale brown(above) to purplish cream (below), immaculate. Concealedsurfaces of arms, hands, shanks, thighs, and feet purple.Anterior and posterior surfaces of thigh with 9–10 darkbrown transverse stripes; superior surfaces reddish brown,stripe in depigmented areas continuous with dark brownstripes on anterior and posterior faces of thigh. Whitelichenose marks dorsal to cloacal opening and heels. Throat,chest, and belly cream, the latter somewhat reddish on distalparts. Iris gold, with outer black aureole. Pupil black.Daytime coloration unknown.

Variation.—Dorsal ground color varies from reddish brownto light brown at night, and become pale cream in daytime.Dorsum immaculate (n 5 26, 58%), with weak dark browntransverse bands on sacral region (n 5 17, 38%; MNRJ25587–88, 29175–76, 29181, 29184, 36940–45, MZUEFS2880, UFBA 6415, 6425, 7485, 7489, 7990), with thin

reticulated brown stripes (MNRJ 36888), or scattered withlarge brown circular markings (MNRJ 29184). Dorsal darkbrown blotches absent (n 5 36, 80%) or present (n 5 9, 20%;MNRJ 29177, 29183, MZUEFS 1269, UFBA 6466, 7000, 7485,7487–88, 7492). Vertebral stripe varies in length. Lichenosespots vary in number and position, including numerous andwidely scattered over dorsum, present on elbows, and absentover cloacal opening and heels. Transverse brown stripes onshanks present (n 5 38, 84%) or absent (n 5 7, 16%; MNRJ29178, 29181, 36888–89, UFBA 7000, 7484, 7491). Trans-verse brown stripes complete (n 5 27, 60%) or partiallyfragmented on posterior surface of thighs (n 5 18, 40%;MNRJ 29176, 29178–79, 36888, 36940, 36942, 36945,MZUEFS 354–55, 1269, 2879, UFBA 6465–66, 7000, 7483,7485, 7488–89). Thin brown transverse bands on flankspresent (n 5 32, 71%) or absent (n 5 13, 29%; MNRJ 25588,36888, MZUEFS 2880, UFBA 6415, 6425, 6437, 6465, 6466,7483, 7485, 7489, 7492, 7990). Transverse brown stripes onthighs 8–11. Two females from the Municipality of Camamu(MNRJ 29183–84) large (SVL 59.8–63.3 mm). Fragmentedpattern of transverse brown bands on dorsum (MNRJ29184), and wider transverse bands on shanks and thighs,also observed in male from the Municipality of Urucuca(MNRJ 25588). Some transverse bands on posterior face ofthighs of males from Amargosa (UFBA 6425), Urucuca(MNRJ 25589), and Wenceslau Guimaraes (MNRJ 29181)

Fig. 3. Holotype of Bokermannohyla capra in life (UFBA 7492), adult male.

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slightly dichotomized. General body proportions similaramong sampled geographic localities (Table 1). Adult maleshave hypertrophied forearms, enlarged prepollices, andvocal slits, characteristics absent in females. Females usuallylarger than males.

Advertisement call.—Advertisement calls varied across geo-graphic samples (Table 2). Social interactions among maleswere evident in variation in structural and temporalparameters of the call. We identified four types of noteswithin advertisement calls of B. capra. The most frequentcall (primary call) comprises only one multi-pulsed note(note-type A; Fig. 4A, B), with pulses grouped within two tonine pulse groups (XX~4:82, n 5 627 notes, SD 5 1.15). Thepulse rate was high enough that it impeded accuratemeasurement of intervals between contiguous pulses,therefore we discarded this temporal parameter. The pulsenumber of one note-type A varied from 30 to 110 pulses(XX~67:68, n 5 523 notes, SD 5 14.13), and is correlatedwith note duration (r2 5 0.30, n 5 453 notes, P , 0.0001);call (5note) duration is 0.24–0.85 s (XX~0:42, n 5 627 notes,SD 5 0.08); inter-note duration is 0.14–15.47 s (XX~1:80, n 5

577 notes, SD 5 1.98); note pulse rate is 88.27–260.50pulses/s (XX~152:68, n 5 453 notes, SD 5 28.97); noterepetition rate is 1.17–4.14 notes/s (XX~2:45, n 5 627 notes,SD 5 0.48). Excited males released the shortest type A notes(0.25–0.53, XX~0:36, n 5 92 notes, SD 5 0.06) and thehighest number of pulse groups (4–9, XX~5:51, n 5 92 notes,SD 5 0.86). This behavior was also noted when one

specimen of B. capra responded to advertisement calls ofAplastodiscus ibirapitanga. Dominant frequencies rangedfrom 0.82 to 2.49 kHz (XX~1:96, n 5 596 notes, SD 5 0.22)within two main energetic bandwidths distinguishable inthe audiospectrogram of type A note (Fig. 4A). Frequencyamplitude ranged from 0.73–2.92 kHz. Alteration in thedominant frequency was observed mainly by a changebetween the energy peaks of intensity in the calls of onespecimen from Camamu.

When producing a series of successive notes of type A,calling males may put more energy in the first released note,which is comprised by only one to two pulse groups, andwith frequency modulation, leading to a clearly distinguish-able initial note (note-type B; Fig. 4A, B). The pulse numberof one note-type B is 28–127 pulses (XX~72:50, n 5 22 notes,SD 5 26.94); note duration is 0.22–0.39 s (XX~0:32, n 5 22notes, SD 5 0.05); interval between a note-type B andanother successive note is 0.16–3.72 s (XX~0:70, n 5 22notes, SD 5 0.71); dominant frequency is 1.80–2.32 kHz(XX~2:01, n 5 22 notes, SD 5 0.14); frequency amplitude is0.99–2.84 kHz.

During duets involving conspecific males, B. capra usuallyproduced a single call, possibly a territorial call. This callcomprised one to four distinct notes (note-types A–E). Thenote-type C resembles note-type A, but presents moreenergy, well-marked pulse groups, and crescent energypeaks from first to last pulse group (Fig. 4C, D). Note-typeC is characterized by 35–85 pulses (XX~59:31, n 5 16 notes,SD 5 13.31), 3–7 pulse groups (XX~4:75, n 5 16 notes, SD 5

Table 1. Descriptive Statistics of Adult Males and Females from the Type-Series of Bokermannohyla capra, New Species, from Serra do Timbo (Type-Locality, Municipality of Amargosa), and from the Combined Samples of Camamu, Elısio Medrado, Itacare, Itubera, Nilo Pecanha, Serra da Jiboia,Urucuca, Valenca, and Wenceslau Guimaraes, All in the State of Bahia, Brazil. n 5 number of specimens; mean 6 standard deviation (range). Otherabbreviations are defined in the text. Measurements are in millimeters.

Males Females

Serra do Timbo Other localities Serra do Timbo Other localitiesn = 14 n = 24 n = 2 n = 5

SVL 51.4 6 2.3 (46.8–54.2) 52.4 6 2.2 (48.3–56.8) 61.5 6 3.7 (58.9–64.1) 61.2 6 2.0 (59.6–63.3)HL 18.5 6 1.0 (16.6–20.7) 19.2 6 0.9 (17.4–21.8) 21.3 6 1.1 (20.5–22.1) 22.4 6 0.5 (21.5–22.8)HW 17.8 6 0.9 (16.0–18.9) 18.5 6 0.8 (17.0–20.4) 21.0 6 1.9 (19.6–22.3) 21.5 6 0.7 (20.7–22.3)ED 5.4 6 0.4 (4.7–6.1) 5.8 6 0.4 (5.0–6.6) 5.9 6 0.4 (5.6–6.1) 6.8 6 0.3 (6.3–6.9)END 5.0 6 0.2 (4.6–5.4) 5.2 6 0.3 (4.8–6.0) 5.7 6 0.3 (5.4–5.9) 6.4 6 0.2 (6.1–6.6)TD 3.0 6 0.2 (2.7–3.4) 3.2 6 0.2 (2.9–3.8) 3.5 6 0.1 (3.4–3.6) 3.7 6 0.1 (3.6–3.7)UEW 4.3 6 0.4 (3.6–4.9) 4.5 6 0.4 (3.6–5.2) 4.5 6 1.2 (3.6–5.3) 5.3 6 0.1 (5.0–5.3)IOD 6.0 6 0.5 (5.0–6.9) 6.0 6 0.3 (5.5–6.4) 6.7 6 0.3 (6.4–6.9) 7.2 6 0.3 (6.9–7.4)IND 3.4 6 0.3 (3.0–4.0) 3.6 6 0.2 (3.3–3.9) 3.8 6 0.3 (3.6–4.0) 4.0 6 0.1 (3.8–4.1)NSD 3.0 6 0.2 (2.8–3.3) 3.0 6 0.2 (2.6–3.3) 3.5 6 0.0 (3.4–3.4) 3.2 6 0.1 (3.0–3.3)THL 26.1 6 1.3 (23.2–27.9) 26.8 61.3 (24.0–28.7) 29.9 6 1.5 (28.9–31.0) 30.3 6 0.7 (29.6–31.0)TL 26.1 6 1.5 (22.6–28.2) 27.2 6 1.3 (24.0–29.3) 29.6 6 4.2 (26.6–32.6) 31.1 6 0.8 (30.4–31.9)FL 35.8 6 1.7 (32.1–38.0) 36.9 61.7 (32.2–39.2) 42.2 6 3.4 (39.8–44.6) 42.3 6 1.5 (41.1–44.0)3FD 2.8 6 0.2 (2.4–3.1) 2.8 6 0.2 (2.2–3.1) 3.2 6 0.2 (3.0–3.3) 3.3 6 0.2 (3.0–3.4)4TD 2.6 6 0.2 (2.4–3.1) 2.6 6 0.2 (2.2–3.1) 3.0 6 0.2 (2.8–3.1) 3.0 6 0.2 (2.7–3.1)SVL/HW 2.89 6 0.05 (2.80–3.01) 2.83 6 0.06 (2.72–2.94) 2.93 6 0.09 (2.87–3.00) 2.84 6 0.01 (2.83–2.87)SVL/HL 2.77 6 0.07 (2.57–2.87) 2.73 6 0.09 (2.53–3.06) 2.88 6 0.01 (2.87–2.90) 2.73 6 0.05 (2.66–2.77)HL/SVL 0.34 6 0.01 (0.33–0.36) 0.37 6 0.01 (0.33–0.39) 0.34 6 0.00 (0.34–0.34) 0.36 6 0.00 (0.36–0.37)ED/TD 1.79 6 0.12 (1.58–2.00) 1.81 6 0.13 (1.50–2.00) 1.67 6 0.06 (1.62–1.72) 1.82 6 0.03 (1.76–1.84)TD/SVL 0.05 6 0.00 (0.05–0.06) 0.06 6 0.00 (0.06–0.06) 0.05 6 0.00 (0.05–0.06) 0.06 6 0.00 (0.05–0.06)ED/SVL 0.10 6 0.00 (0.09–0.11) 0.11 6 0.00 (0.10–0.12) 0.09 6 0.00 (0.09–0.09) 0.11 6 0.00 (0.10–0.11)THL/SVL 0.50 6 0.00 (0.48–0.51) 0.51 6 0.02 (0.46–0.55) 0.48 6 0.00 (0.48–0.49) 0.49 6 0.00 (0.48–0.51)TL/SVL 0.50 6 0.01 (0.48–0.52) 0.52 6 0.02 (0.50–0.56) 0.48 6 0.04 (0.45–0.50) 0.50 6 0.00 (0.50–0.51)THL+TL 52.25 6 2.75 (45.8–56.1) 53.96 6 2.53 (48.0–58.0) 59.55 6 5.72 (55.5–63.6) 61.40 6 1.45 (60.0–62.9)

Napoli and Pimenta—New Bokermannohyla from Bahia, Brazil 679

1.23), and 0.25–0.65 s (XX~0:42, n 5 16 notes, SD 5 0.12)note duration; intervals between a note-type C and anothersuccessive note vary from 0.11–2.12 s (XX~0:32, n 5 16notes, SD 5 0.52). The dominant frequency is 1.80–2.32 kHz(XX~2:01, n 5 16 notes, SD 5 0.24) and frequency amplitudeis 1.03–2.97 kHz. Note-type C is always followed by one tofour short notes, the note-type D. The pulse number of onenote-type D is 8–23 pulses (XX~15:76, n 5 34 notes, SD 5

3.62), 1–2 (XX~1:11, n 5 34 notes, SD 5 0.32) pulse groups,and 0.06–0.15 s (XX~0:09, n 5 34 notes, SD 5 0.02) noteduration. The interval between a note-type D and anothersuccessive note is 0.07–9.74 s (XX~1:72, n 5 34 notes, SD 5

2.57); dominant frequency is 1.80–2.15 kHz (XX~1:96, n 5

22 notes, SD 5 0.12); frequency amplitude is 0.94–2.84 kHz.Note-type D is always followed by a short to very short note(note-type E). Note-type E is similar to note-type C indominant frequency (1.80–2.32 kHz, XX~2:05, n 5 14 notes,SD 5 0.21) and in having well-marked pulse groups (1–5,XX~3:2, n 5 19 notes, SD 5 0.85); it differs from the lattermainly by lacking the first pulse groups, which leads to ashorter duration (note duration 0.12–0.43 s, XX~0:24, n 5 19

Fig. 4. Advertisement call of Bokermannohyla capra. Primary call:audiospectrogram (A) and oscillogram (B) of note-types A and Breleased by the holotype (UFBA 7492). Territorial call: audiospectro-gram (C) and oscillogram (D) of note-types C, D, and E produced by amale from the paratopotypes UFBA 7483–88, 7490–91 (the identifi-cation tag of the calling male was lost during the fieldwork).Vocalizations recorded on 18–19 July 2007; air temperature 21.5uC;air moisture 74%.

Tab

le2.

Ad

vert

isem

en

tC

allP

ara

mete

rso

fN

ote

-Typ

eA

(Pri

mary

Call)

ofB

oke

rma

nnoh

yla

cap

ra,N

ew

Speci

es,

per

Geo

graph

icSam

ple

inB

ahia

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te,B

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l.N

um

ber

of

pu

lses

an

dp

uls

ere

petition

rate

were

no

tm

easu

red

for

Cam

am

usp

eci

men

sd

ue

toexc

ess

ive

en

viro

nm

enta

ln

ois

ew

ithin

no

tes.

Cal

ld

ura

tio

nan

din

terc

all

inte

rvalin

seco

nd

s;fr

eq

uen

cies

inkH

z.

Am

arg

osa

Cam

am

ua,b

Vale

nca

W.G

uim

ara

esb

Anal

yzed

calls

/mal

e4

53

/61

48

/26

0/1

25

/1Puls

egr

oups/

call

2–

9(4

.76

1.1

)3

–7

(4.6

61

.03

)3

–6

(4.7

60

.7)

6–

9(6

.96

0.8

8)

Puls

es/c

all

39

–1

10

(69

.06

14

.3)

—3

0–

78

(58

.66

9.3

)4

1–

87

(61

.26

10

.2)

Note

dura

tion

(s)

0.2

7–

0.8

5(0

.45

60

.07

)0

.24

–0

.40

(0.3

26

0.0

3)

0.2

4–

0.4

4(0

.35

60

.04

)0

.32

–0

.53

(0.4

56

0.0

5)

Inte

r-note

dura

tion

(s)

0.1

9–

15

.47

(1.9

66

1.9

4)

0.1

7–

3.8

5(0

.73

60

.59

)0

.17

–1

1.1

0(3

.45

60

.05

)0

.14

–0

.50

(0.3

06

0.0

7)

Puls

era

te(p

uls

es/s

)88

.2–

26

0.5

(15

2.9

629

.69

)—

108.2

–253.1

(161.8

622.2

)1

11

.7–

19

6.3

(13

2.4

61

6.1

)N

ote

rate

(note

s/s)

1.1

7–

4.4

8(2

.28

60

.37

)2

.45

–4

.01

(3.1

06

0.3

2)

2.2

6–

4.1

4(2

.85

60

.41

)1

.85

–3

.04

(2.2

36

0.3

1)

Dom

inan

tfreq

uen

cy1

.80

–2

.49

(2.0

56

0.1

3)

0.8

2–

2.0

6(1

.75

60

.19

)1

.17

–2

.39

(1.8

36

0.3

7)

1.9

5–

2.1

0(2

.01

60

.04

)Fr

equen

cyam

plit

ude

0.7

3–

2.9

20

.47

–2

.00

1.1

7–

2.3

91

.07

–2

.32

aD

om

inan

tfreq

uen

cyof

one

reco

rded

spec

imen

varie

dbet

wee

ntw

ohar

monic

s(c

a.0

.9an

d1

.8kH

z,re

spec

tivel

y)b

Exci

ted

spec

imen

s

680 Copeia 2009, No. 4

notes, SD 5 0.07), and by the decrease of energy peaks fromfirst to last pulse group.

The currently known advertisement calls of other speciesof the B. circumdata group are markedly different in overallstructure from B. capra. Assuming that we are comparinghomologous calls (primary notes released in the respectiveadvertisement calls), the advertisement call of B. capra mostresembles the calls of B. hylax and B. lucianae, differing fromboth by the number of pulses and overall structure of thecall. From B. hylax, the call of B. capra is differentiated by thepresence of nearly superimposed pulse groups in the firsthalf of the note, with wider intervals between pulse groupsin the end of the note (in B. hylax, pulse groups are wideapart from each other initially, and nearly superimposed inthe end of the note); also, in B. capra note-types A and Bexhibit frequency modulation (absent in B. hylax). From B.lucianae, the new species differs in having more regularlydistributed pulses, without well-defined pulse groups in theprimary call (note-type A), greater number of pulses per note(B. capra: 30–110 pulses/note, XX~67:68, n 5 523 notes, SD 5

14.13; B. lucianae: 15–37, XX~27:60, n 5 48 notes, SD 5

4.89), and greater pulse rate (B. capra: 88.27–260.50 pulses/sec, XX~152:68, n 5 453 notes, SD 5 28.97; B. lucianae:54.81–93.33, XX~72:47, n 5 48 notes, SD 5 9.24).

Natural history.—Bokermannohyla capra is restricted to Atlan-tic Forest remnants. At Serra do Timbo, in the municipalityof Amargosa, specimens were widespread, calling in theforest, often on bushes near forest brooks, up to ca. 3 mfrom the ground. Some were calling from cavities in treebark and bromeliads. Reproductive activity was observedduring all seasons. Tadpoles were captured at night undersubmersed leaf litter from a shallow forest brook (ca. 5 cmdepth) in which calling males were observed, but were notadequately preserved to permit description of larvae. Junca(2006) reported that B. hylax (5B. capra) was among themost abundant anurans at Serra da Jiboia, inhabitingpermanent and temporary water bodies. Specimens fromCamamu and Wenceslau Guimaraes were collected insidesecondary or well-preserved Atlantic Forest fragments. Thesefragments had in common a well-structured canopy, diversetypes of aquatic habitats, and abundance of epiphytebromeliads. Males were seen calling from bromeliads up toeight meters from the ground on trees near water, perchedon marginal vegetation of small streams, or from thevegetation of floodplains. Females were captured at nighton vegetation near a temporary pond.

Geographic distribution.—Bokermannohyla capra is knownonly from coastal Atlantic Forest remnants in eastern Bahia,northeastern Brazil, within regions known as Vale doJiquirica, Baixo Sul, and Litoral Sul. The type locality, Serrado Timbo, is a mountain characterized by rugged reliefcovered mainly by tropical semi-deciduous forests, withelevations ranging from 800–900 m. The Serra do Timbo lieswithin the municipalities of Amargosa, Ubaıra, and Brejoes,in the State of Bahia, Brazil, and belongs, together with theSerra da Jiboia (12u529S, 39u289W), to a set of disjunctmountains located north of the Serra Geral mountain range.The Serra do Timbo is in the western part of the geographicregion known as Reconcavo Baiano, and is approximately95 km northeast from the Municipality of Maracas (13u269S,40u259W). This region was considered by Ab’Saber (1977) tobe a transitional area between the Caatinga, Cerrado, and

Tropical Atlantic domains. The tropical-humid climateprevails in the Serra do Timbo, but the region is surroundedby sub-humid and semi-arid climates.

Paratypes collected outside the type locality and Serra daJiboia were all found in tropical rainforests of southernBahia (Baixo Sul and Litoral Sul), most located in low coastalplains (except for Urucuca and Wenceslau Guimaraes thatinclude mountains with elevations up to 900 m). These arecharacterized by tropical-humid climate, with rains occur-ring during all months of the year (see Jardim [2003] fordetailed descriptions on forest structure of most of the areaswhere paratypes were found).

Bokermannohyla capra is not sympatric with any otherBokermannohyla and represents the northernmost distribu-tion limits of the B. circumdata species group in the AtlanticForest Domain (Fig. 5), with all known geographic samplesoccurring north of the Cachoeira river basin. This basin isplaced in southeastern Bahia, and includes the Colonia andCachoeira as its main rivers, the latter with its mouth in theMunicipality of Ilheus.

Bokermannohyla hylax, one of the two species that mostresembles B. capra morphologically, is distributed withinmountain ranges in southern and southeastern Brazil (Serrado Mar and Serra da Mantiqueira mountain ranges), and isseparated from B. capra by a geographic distance of approx-imately 1000 km. Bokermannohyla lucianae, the species thatmost resembles B. capra, is known only from the Municipalityof Una (State of Bahia), south of the Cachoeira river basin.

Conservation.—Three sampled populations of B. capra occurin two different kinds of protected areas: at the State level(Estacao Ecologica Estadual Nova Esperanca, Municipality ofWenceslau Guimaraes), or at the Private level (ReservaMichelin, Municipality of Itubera; RPPN Agua Branca,Municipality of Valenca). On the other hand, at the typelocality, the forest cover of Serra do Timbo and its associatedbiota are currently threatened by human activities, mainlyby deforestation for the development of banana and cacaocultivations, pastures, and selective logging of species ofcommercial interest. Concomitantly, the springs that rise inSerra do Timbo and support the water supply of themunicipalities of Amargosa, Ubaıra, Matuıpe, and Jiquiricaare also endangered. Currently, the Serra do Timbo is theobject of a conservation movement (Timbo’s Project) thatseeks to protect natural resources including plant andanimal species as well as their habitat. The Timbo’s Projectis developed by the Centro Sapucaia, Nucleo Mata Atlantica(Ministerio Publico), and by the municipal government ofAmargosa. This action includes the creation of a conserva-tion unit with 674 ha of tropical semi-deciduous forest,which will be supported by the Ministerio do MeioAmbiente (data obtained from Centro-Sapucaia, 2008,accessible at http://www.centrosapucaia.org.br).

Localities on Baixo Sul and Litoral Sul are all threatenedmainly by deforestation, which still occurs regularly despiteprohibition even in legally protected areas. The conversionof forests to banana, coffee, eucalyptus in more recent times,and other cultivars is also contributing to habitat loss insouthern Bahia. This state is now experiencing a new boostof infrastructure development, including road and railroadconstruction and mining. Their impacts on the fragileenvironments of the Mata Atlantica are worthy of concern.Many projects and enforcement actions are being conductedby governmental and civil authorities and other agencies to

Napoli and Pimenta—New Bokermannohyla from Bahia, Brazil 681

avoid deforestation in the region, including the creation ofnew reserves.

Etymology.—The specific name, a Latin substantive (caper 5

goat), is an allusion to the advertisement call of the newspecies, which sounds like a bleating of a goat.

ACKNOWLEDGMENTS

We are grateful to C. Cruz for critically reading andimproving the manuscript; F. Junca and U. Caramaschi forallowing us to examine specimens under their care; theCentro de Desenvolvimento Sustentavel e AgroecologiaSapucaia–Centro Sapucaia, M. Neves, and R. Neto forallowing us to access research areas under their care andfor logistical assistance at Serra do Timbo; Superintendenciade Desenvolvimento Florestal e Unidades de Conservacaofor the permission to work at Estacao Ecologica EstadualNova Esperanca and for logistical assistance, which wereprovided by M. Silva and E. Santos (Nego); the owners ofFazenda Caititu, Fazenda Sao Joao, Projeto de Assentamento

Zumbi dos Palmares (especially J. Silivrano), and RPPN AguaBranca for permission to work on their properties; themunicipal government of Amargosa for logistical support; R.Abreu for line drawings and photographs of the holotype; P.Cordeiro, P. Fonseca, R. Abreu, R. Lopes, T. Silva, and M.Freitas for help during field work; G. Woehl, Jr. (Instituto Ra-bugio para Conservacao da Biodiversidade) for allowing usto examine advertisement calls of B. hylax from theMunicipality of Guaramirim, State of Santa Catarina, Brazil.Most collections on Baixo Sul and Litoral Sul were madeduring the sub-project ‘‘Abordagens Ecologicas e Instrumen-tos Economicos para o Estabelecimento do Corredor doDescobrimento: uma Estrategia para Reverter a Fragmenta-cao Florestal da Mata Atlantica no Sul da Bahia,’’ supportedby the Projeto de Conservacao e Utilizacao Sustentavel daDiversidade Biologica Brasileira/Ministerio do Meio Am-biente (PROBIO/MMA), Center for Applied BiodiversityScience/Conservation International (CABS/CI), World Bank,and coordinated by the Instituto de Estudos Socio-Ambien-tais do Sul da Bahia (IESB), and also during the Project‘‘Planejamento Participativo em RPPN no Corredor Central

Fig. 5. Geographic distribution of Bokermannohyla capra and most related species on a topographic map. Geographic distribution of B. hylax basedon Napoli (2000). Numbered circles represent main hydrographic basins in southeastern Bahia (only major rivers are indicated): 1, Contas riverbasin; 2, Cachoeira river basin; 3, Pardo river basin; 4, Jequitinhonha river basin.

682 Copeia 2009, No. 4

da Mata Atlantica,’’ funded by the Fundo Nacional de MeioAmbiente/Ministerio do Meio Ambiente (FNMA/MMA) andcoordinated by the Instituto de Estudos Socio-Ambientais doSul da Bahia (IESB). The Instituto Brasileiro do MeioAmbiente e dos Recursos Naturais Renovaveis (IBAMA)permitted collection of specimens.

LITERATURE CITED

Ab’Saber, A. N. 1977. Os domınios morfoclimaticos naAmerica do Sul. Primeira aproximacao. Geomorfologia52:1–23.

Bokermann, W. C. A., and I. Sazima. 1973. Anfıbios daSerra do Cipo, Minas Gerais, Brasil. 1. Especies novas de‘‘Hyla’’ (Anura, Hylidae). Revista Brasileira de Biologia33:329–336.

Caramaschi, U., M. F. Napoli, and A. T. Bernardes. 2001.Nova especie do grupo de Hyla circumdata (Cope, 1870) doEstado de Minas Gerais, Brasil (Amphibia, Anura, Hyli-dae). Boletim do Museu Nacional, Nova Serie, Zoologia457:1–11.

Cardoso, A. J., and G. V. Andrade. 1982. Nova especie deHyla do Parque Nacional Serra da Canastra (Anura,Hylidae). Revista Brasileira de Biologia 42:589–593.

Duellman, W. E., and L. Trueb. 1994. Biology of Amphib-ians. Second edition. John Hopkins University Press,Baltimore and London.

Faivovich, J., C. F. B. Haddad, P. C. A. Garcia, D. R. Frost,J. A. Campbell, and W. C. Wheeler. 2005. Systematicreview of the frog family Hylidae, with special reference toHylinae: phylogenetic analysis and taxonomic revision.Bulletin of the American Museum of Natural History294:1–240.

Heyer, W. R., A. S. Rand, C. A. G. Cruz, O. L. Peixoto, andC. E. Nelson. 1990. Frogs of Boraceia. Arquivos deZoologia 31:231–410.

Jardim, J. G. 2003. Uma caracterizacao parcial da vegetacaona regiao sul da Bahia, Brasil. In: Corredor de Biodiversi-dade na Mata Atlantica do Sul da Bahia. P. I. Prado, E. C.

Landau, R. T. Moura, L. P. S. Pinto, G. A. B. Fonseca, andK. N. Alger (eds.). IESB/CI/CABS/UFMG/UNICAMP, Il-heus, Brazil, CD-ROM.

Junca, F. A. 2006. Diversidade e uso de habitat por anfıbiosanuros em duas localidades de Mata Atlantica, no norte doestado da Bahia. Biota Neotropica 6:1–17.

Myers, C. W., and W. E. Duellman. 1982. A new species ofHyla from Cerro Colorado, and other tree frog records andgeographical notes from Western Panama. AmericanMuseum Novitates 2752:1–32.

Napoli, M. F. 2000. Taxonomia, variacao morfologica edistribuicao geografica das especies do grupo de Hylacircumdata (Cope, 1870) (Amphibia, Anura, Hylidae).Unpubl. Dr.Sc. diss. Universidade Federal do Rio deJaneiro/Museu Nacional, Rio de Janeiro, Brazil.

Napoli, M. F. 2005. A new species allied to Hyla circumdata(Anura: Hylidae) from Serra da Mantiqueira, SoutheasternBrazil. Herpetologica 61:63–69.

Napoli, M. F., and U. Caramaschi. 2004. Two new speciesof the Hyla circumdata group from Serra do Mar and Serrada Mantiqueira, Southeastern Brazil, with description ofthe advertisement call of Hyla ibitipoca (Anura, Hylidae).Copeia 2004:534–545.

Napoli, M. F., and F. A. Junca. 2006. A new species of theBokermannohyla circumdata group (Amphibia: Anura: Hy-lidae) from Chapada Diamantina, State of Bahia, Brazil.Zootaxa 1244:57–68.

Napoli, M. F., and B. V. S. Pimenta. 2003. Nova especie dogrupo de Hyla circumdata (Cope, 1870) do sul da Bahia,Brasil (Amphibia, Anura, Hylidae). Arquivos do MuseuNacional 61:189–194.

Pombal, J. P., and C. F. B. Haddad. 1993. Hyla luctuosa, anew treefrog from southeastern Brazil (Amphibia, Hyli-dae). Herpetologica 49:16–21.

Savage, J. M., and W. R. Heyer. 1967. Variation anddistribution in the tree-frog genus Phyllomedusa in CostaRica, Central America. Beitrage zur Neotropischen Fauna5:111–131.

Napoli and Pimenta—New Bokermannohyla from Bahia, Brazil 683