A new species of Trachycephalus (Amphibia, Anura, Hylidae) from the State of Goiás, Brazil

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Accepted by M. Vences: 23 Oct. 2009; published: 12 Jan. 2009 58

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A new species of Trachycephalus (Amphibia, Anura, Hylidae) from the State of Goiás, Brazil

CARLOS EDUARDO DOMINGOS CINTRA1, HÉLDER LÚCIO RODRIGUES SILVA1,2,

NELSON JORGE DA SILVA JR.1,2,3, PAULO CHRISTIANO DE ANCHIETTA GARCIA4,5,

& HUSSAM ZAHER6

1Systema Naturae Consultoria Ambiental Ltda. Rua 58 no217 – Jardim Goiás. CEP 74810-250 – Goiânia, GO. BRAZIL2Centro de Estudos e Pesquisas Biológicas. Universidade Católica de Goiás, Avenida Universitária, 1440 – Setor Universitário. CEP 74605-010 – Goiânia, GO. BRAZIL3Mestrado em Ciências Ambientais e Saúde. Universidade Católica de Goiás, Rua 232, no 128 – 3º andar – Área V – Setor Universi-tário. CEP 74605-140 – Goiânia, GO. BRAZIL4Museu de Zoologia da Universidade de São Paulo. Avenida Nazaré 481 – Ipiranga. CEP 04263-000 – São Paulo, SP. BRAZIL. E-mail: [email protected] de Zoologia, ICB, Universidade Federal de Minas Gerais, Avenida Antônio Carlos, 6627 – Pampulha – CEP 31270-901 - Belo Horizonte, MG. BRAZIL6Corresponding author

Abstract

A new species of Trachycephalus is described for the Cerrado biome of Goiás, Brazil. Trachycephalus mambaiensis sp.nov. is distinguished from the other ten species of the genus by the skin co-ossified with the skull, heavy cranial ossifica-tion, frontoparietal that fails to articulate with squamosal, absence of a crista occipitalis and secreting glands of milkyand viscous substances. The skull of the new species shows an intermediary condition between species of Trachycepha-lus with a well ossified skull (Casque-headed frogs) and those without cranial ossification.

Key words: Cerrado, taxonomy, Trachycephalus, new species, osteology

Resumo

Uma nova espécie de Trachycephalus é descrita do Bioma Cerrado de Goiás, Brasil. Trachycephalus mambaiensis sp.nov. caracteriza-se das outras dez espécies do gênero por apresentar a pele da cabeça aderida ao crânio, grande ossifica-ção cranial e ausência de crista occipital e glândulas secretoras de substancias leitosas e viscosas. Osteologicamente, ocrânio da nova espécie mostra uma condição intermediária entre as espécies de Trachycephalus com crânio ossificado(Casque-headed tree frogs) e as espécies com crânio não ossificado.

Palavras chave: Cerrado, taxonomia; Trachycephalus, nova espécie, osteologia

Introduction

In a recent molecular phylogenetic analysis of Hylid frogs, Faivovich et al. (2005) suggested that the genusPhrynohyas Fitzinger 1843 was paraphyletic in respect to the genus Trachycephalus Tschudi 1838, syn-onymising the former with the latter in order to provide a monophyletic taxonomy for the group. The genusTrachycephalus now comprises ten species of medium-sized arboreal anurans broadly distributed throughoutSouth America (Frost, 2007). The monophyly of the newly defined genus Trachycephalus is well supported


Zootaxa 1975 © 2009 Magnolia Press · 59 NEW SPECIES OF TRACHYCEPHALUS FROM CENTRAL BRAZIL

by 37 synapomorphic protein transformations (Faivovich et al., 2005) and one morphological synapomorphy:a double vocal sac that is laterally disposed and projects posteriorly to the mandible when inflated (Duellman,1968, Trueb and Duellman, 1971).

The ten known species of Trachycephalus are T. atlas Bokermann 1966, T. coriaceus (Peters 1867), T.hadroceps (Duellman and Hoogmoed 1992), T. imitatrix (Miranda-Ribeiro 1926), T. jordani Stejneger andTest 1891, T. lepidus (Pombal, Haddad, and Cruz 2003), T. mesophaeus (Hensel 1867), T. nigromaculatusTschudi 1838, T. resinifictrix (Goeldi 1907), and T. venulosus (Laurenti 1768), from which only the latter twoare known to occur in the Brazilian state of Goiás.

Five specimens of an unknown species of Trachycephalus were collected between July and September2006 in a region of transition between Cerrado and Caatinga biomes, in the municipality of Mambaí, locatedin the Northern part of the State of Goiás. The specimens were compared with representatives of the knownspecies of the genus and revealed a combination of diagnostic characters that allows its description as a newspecies.

Material and methods

We analyzed 107 specimens, representing nine of the 10 species presently known for the genus Trachycepha-lus (Appendix), belonging to the following collections (acronyms in parenthesis): Coleção de Herpetologia daUniversidade de Brasília (CHUNB), and Museu de Zoologia da Universidade de São Paulo (MZUSP). Wealso examined dried skulls from adult specimens of eight species, including the new species (Appendix).

The holotype and paratypes were radiographed with the aid of a Faxitron x-ray, model Mx 20. Adult spec-imens were measured to the nearest 0.01 mm with a digital caliper. Measurements follow Duellman (1970),Cei (1980), and Heyer et al. (1990). Abbreviations used in the measurements are as follow: SVL (snout-ventlength), HL (head length), HW (head width), TD (tympanum diameter), ED (eye diameter), AMD (distancebetween the anterior margins of eyes), EN (eye to nostril distance), ID (internarial distance), NS (nostril to tipof snout distance), FAL (forearm length), FAB (forearm breadth), HAL (hand length), THL (thigh length),TBL (tibia length), TAL (tarsal length), and FL (foot length). Webbing formulae follow Savage and Heyer(1967), as modified by Myers and Duellman (1982).

Trachycephalus mambaiensis sp. nov.Figs. 1–4

Holotype. MZUSP 135715, an adult male (Fig. 1) from the vicinity of Santa Edwiges I hydroelectric powerplant dam, Municipality of Mambaí, State of Goiás, Brazil (14°17'24''S 46°11'37''W or UTM 0371233 and8419860) (Fig. 2), collected by Carlos Eduardo Domingos Cintra on 2 August 2006.

Paratypes. Two adult females (MZUSP 135713, 135716) and two adult males (MZUSP 135714, 135717)(Fig. 3) from the vicinity of the Santa Edwiges I hydroelectric power plant, Municipality of Mambaí, State ofGoiás, collected by Carlos Eduardo Domingos Cintra on 31 July and 17 August 2006 (Fig. 2); two adult males(CHUNB 21868, 35471) from the municipality of São Domingos, State of Goiás, collected by Laurie J. Vittand Janalee P. Caldwell between 19 November and 15 December 2003.

Diagnosis. The species is included in the genus Trachycephalus which present paired vocal sacs protrud-ing posterior to the angles of the jaws when inflated as a unique synapomorphy (fide Trueb and Duellman,1971; Tyler, 1971; Faivovich et al., 2005). A medium-sized species of Trachycephalus (SVL in males76.4–82.0 mm; females 83.67–86.71 mm) characterized by the combination of the following traits: presenceof paired vocal sacs protruding posterior to the angles of the jaws when inflated; a rounded snout without amedial (premaxillary) notch in dorsal view and truncate in lateral view; skin co-ossified with the skull; fron-


CINTRA ET AL.60 · Zootaxa 1975 © 2009 Magnolia Press

toparietals converge medially throughout their lengths, covering the fontanelle; frontoparietals not coveringthe prootics posteriorly; frontoparietals not articulating with squamosals; presence in life of small reddishspots sparsely disposed on the dorsum.

FIGURE 1. Trachycephalus mambaiensis, MZUSP 135715 (holotype), adult male from the vicinity of the Hydroelectricdam Santa Edwiges I, Municipality of Mambaí, State of Goiás, Brazil, (A) dorsal view; (B) Ventral view; (C) head in lat-eral view. Scale = 10 mm.

The new species differs from T. coriaceus, T. hadroceps, T. imitatrix, T. lepidus, T. mesophaeus, T. resin-ifictrix, and T. venulosus by the presence of a well ossified skull in which the skin is adhered, and from T.atlas, T. jordani, and T. nigromaculatus by frontoparietals that fail to cover the prootics posteriorly and by thelack of an occipital crest and medial premaxillary notch. Further, the new species differs from all ten speciesof Trachycephalus (except T. nigromaculatus) by the presence of small reddish spots on the dorsum.

Description of the holotype. Body robust; head narrower than the body, as wide as long (Fig. 1); top ofthe head flat, skin visibly adhered to the skul; snout short, rounded in dorsal profile and nearly truncate in lat-eral profile; canthus rostralis rounded distinct; loreal region barely concave; lips moderately thin and flared;nostrils protuberant, directed laterally; distance of nostril to tip of snout smaller than internarial and eye-nos-tril distances; internarial region slightly depressed; eyes lateral, directed anteriorly, moderate size; eyes diam-eter being close to 1/4 of head length; pupil horizontal; tympanum distinct, large, its diameter approximately



four-fifth of the eye diameter; supratympanic fold extends posteriorly from the posterior corner of the eye,above the tympanum. Vocal sacs paired and lateral, behind the angle of the jaw; vocal slits large, located later-ally under the tongue; tongue large, cordiform, covering approximately 2/3 of the mouth floor, free andnotched posteriorly; dentigerous processes of prevomer slightly curved in a transverse plane, each one bearingeigth and seven teeth. Arms short, moderately robust; without axillary membrane and distinct ulnar fold;hands large, length slightly smaller than head length (Fig. 4A); fingers medium sized and moderately robust;finger discs large; width of the disc on the third finger smaller than the tympanum diameter; subarticulartubercles single, large and oval on finger I-IV; outer palmar tubercle large, bifid; inner palmar tubercle large,elliptical; numerous palmar supranumerary tubercles; finger disks large, nearly rounded; relative fingerlengths: I < II < IV < III; fingers slightly webbed; webbing formula: I – II 2- - 3+ - 2+ IV. Legs short, robust;thighs slightly smaller than mead of SVL; feet relatively small, about 1/3 of SVL; robust toes; toe discs large

(Fig. 4B); relative toe lengths: I < II < V ˜ III < IV; foot webbing formula: I 1 1/2 – 2 +II 1 1/3 – 2 1/2 III 1 1/

2– 3- IV 2 1/2 – 1+ V; inner metatarsal tubercle larger, elliptic, visible from dorsal view; outer metatarsal tuber-cle large, bifid. Toes moderately short; toe discs slightly smaller than those on fingers; subarticular tuberclesmoderately large, subconical; small supernumerary metatarsal tubercles; inner tarsal fold absent. The dorsalskin texture is finely granular; lateral and undersurface skin texture smooth.

Variation. Individuals of Trachycephalus mambaiensis show some variation in size (Table 1), develop-ment of toe, finger webbing, and pattern of dorsal blotches. The holotype is the largest male of the series.Females are slightly larger than males, have larger fingers and toes, more extensive webbing.

Color in life of the holotype. Dorsum dark brown with several irregular light brown or creamy blotches.There are two wide and long, dorsolateral creamy blotches that extend from the scapular to the sacral region;some small, round red-cooper blotches randomly distributed on the dorsum and flanks; several large, roundcreamy blotches on a brown-grayish background on the flanks; dorsal surfaces of the anterior and posteriorlimbs show creamy transversal bars on a dark brown background; dorsum of fingers and toes yellowish; ven-tral surface of the body is creamy, with irregular gray spots disposed mostly in the gular region and flanks. Irisgolden and finely reticulated with black. If not inflated, vocal sacs vary from dark brown to black.

Color in preservative of the holotype. Dorsum brownish-gray with creamy blotches; lighter surfaces ofthe venter and flanks grayish white (small red blotches on the body in life are not noticeable). Dorsal surfacesof fingers and toes without any differential coloration related to the limbs.

Cranial osteology. (based in an adult female paratype MZUSP 135713; Fig. 5) skull of T. mambaiensisslightly wider than long (cranium length 24.22mm; cranium width 25.05mm; cranium height 8.45mm). Snoutin dorsal view truncate and bears a poor medial notch; all dorsal surfaces of the dermal bones sculptured intorugosities and irregular ridges; distal margins of the dermal bones rugose or slightly spinose; premaxillariesrobust and lie anteromedially to the maxillaries; left and right pars dentalis of the premaxillae bear 11 and 13teeth, respectively; alary processes of the premaxillaries slightly expanded and co-ossified anteriorly, slightlyconcave posteriorly and oriented vertically; it approach but not articulate with the anterior tip of the nasalanterodorsally; dorsomedially, the alar processes form part of the anterior margin of the external nares; later-ally, the alary processes approach but not to articulate with the pars facialis of the maxillary, whereas mediallyarticulate with one another; the pars palatina of the premaxillary developed and the posteromedial palatineprocess conspicuous. Maxillary moderately robust, but lacks a labial flange; the pars facialis well developed,it is larger anteriorly to the orbit, and articulates with the alary process of the premaxillary to form the ventralmargin of the external naris; posterior to the external naris, the dorsal edge of the pars facialis is free, andarticulates with the maxillary process of the nasal at the anterior margin of the orbit; the pars facialis dimin-ishes in size ventrally to the orbit, but is conspicuous along the entire length of the maxillary; the pars palatinaof the maxillary well developed and extends from the anterior end of the bone posteriorly to the level of termi-nation of the pars dentalis; posteriorly, the maxillary firmly articulates with the quadratojugal which lies medi-ally, adjacent to the maxillary; the outer surface of the maxillary is co-ossified and slightly rugose, principally



in the anterior part; each maxillae bears approximately 70 pedicillate teeth. Quadratojugal increases in sizeposteriorly, whereas the maxillary diminishes in size; posteriorly, ossification of the quadratojugal invades thecartilage of the quadrate process.

TABLE 1. Measurements (mm) and body ratios (%) of Trachycephalus mambaiensis: , arithmetic mean; SD, standarddeviation.

In ventral view, the paired vomer lies lateral to the midline of the skull and ventral to the sphenethmoid;anterior end of vomer contacts the premaxillary-maxillary suture dorsolaterally; vomer bears well developedlateral wings which form the anterior and medial margins of the internal naris; vomers converge medially justanterior to the vomerine teeth; the dentigerous processes slightly curved in a transverse plane, each one bear-ing nine teeth; vomer not contact the neopalatines; paired neopalatine long, robust, and bears a ventral ridge

Measurements HolotypeMZUSP 135715

Males including the Holotype (n = 7)

± SD (Range)

Females(n = 2)

SVL 82.03 78.64 ± 2.43 (76.4–82.0) 83.67–86.71

HL 24.84 24.52 ± 0.86 (23.1–25.4) 28.17–29.19

HW 24.8 24.47 ± 0.95 (23.1–25.7) 26.83–27.8

ED 6.43 6.55 ± 0.46 (6.0–7.1) 7.21–7.47

TD 5.35 5.25 ± 0.15 (5.0–5.4) 5.92–6.14

END 6.98 6.84 ± 0.61 (6.1–7.8) 8.50–8.81

NSD 3.32 3.16 ± 0.18 (2.9–3.3) 3.47–3.35

IND 5.13 4.96 ± 0.20 (4.7–5.1) 5.13–5.32

IOD 10.39 10.43 ± 0.56 (9.7–11.1) 11.22–11.63

FAL 12.19 12.35 ± 0.80 (11.4–13.4) 14.05–16.87

HAL 23.7 23.43 ± 0.84 (22.1–24.2) 25.35–26.27

THL 35.76 35.30 ± 1.36 (33.3–37.1) 40.04–41.68

TL 35.61 35.22 ± 1.39 (33.2–37.0) 39.9–42.22

TAL 19.45 18.88 ± 0.61 (18.1–19.5) 20.25–21.01

FL 30.65 30.44 ± 0.58 (29.5–31.0) 33.93–34.33

Ratios Holotype Range Range

HL/SVL 30.3% 30.3%–32.6% 33.7%–34.4%

HW/SVL 30.2% 30.2%–33.6% 32.1%–32.5%

HAL/SVL 28.9% 28.9%–31.5% 30.3%–30.3%

THL/SVL 43.6% 43.6%–48.3% 48.1%–48.7%

TL/SVL 43.4% 43.4%–48.3% 48.7%–48.5%

TAL/SVL 23.7% 23.6%–25.3% 24.2%–24.6%

FL/SVL 37.4% 36.7%–40.6% 39.6%–41.3%

HW/HL 99.8% 96.3%–103.2% 95.2%–94.3%

ED/HL 25.9% 25.9%–27.9% 25.6%–25.0%

TD/HL 21.5% 21.1%–21.5% 21.0%–20.6%

END/HL 28.1% 24.1%–31.1% 30.2%–29.5%

NSD/HL 13.4% 11.4%–13.4% 11.9%–11.6%

IND/HL 20.7% 18.9%–20.7% 18.2%–17.8%

IOD/HL 54.1% 54.1%–55.4% 54.9%–53.7%

TD/ED 83.2% 75.8%–83.2% 82.2%–82.2%



FIGURE 2. Type locality of Trachycephalus mambaiensis, Hydroelectric dam Santa Edwiges I, Municipality of Mam-baí, State of Goiás, Brazil (14°17'24''S 46°11'37''W or UTM 0371233 and 8419860).

without odontoids; the distal end of the neopalatine embedded in connective tissue and lies dorsally to the parspalatina of the maxillary and adjacent to the anterior maxillary process; proximal end of the neopalatinepointed and lies ventral to the sphenethmoid and posterior to the prevomer, without a medial contact; paras-phenoid robust and tetraradiate; acuminate anterior end lies slightly posterior to the level of the palatines, ven-trally to the sphenethmoid; lateral alae approximately half the length of the cultriform process; lateral alaeinvest medially to the prootic; posteriorly, parasphenoid extends to the edge of the foramen magnum. In dorsalview, nasals large; anterior end of the nasal blunt and articulates with the dorsal margin of the alary process ofthe premaxilla; laterally, the nasal arches over the external nares and articulates posteriorly and ventrolaterallywith the pars facialis of the maxillary at the corner of the orbit; the nasal forms the anterior margin of theorbit; articulates with the frontoparietal dorsolaterally and the dorsally exposed dermal sphenethmoid dorso-medially; the nasals converge medially anterior to the dermal sphenethmoid; canthal ridge distinct, not pro-nounced; it extends from the anterior end of the nasal along the lateral margin to the anterodorsal corner of theorbit; entire nasal surface involved in the integumentary-cranial co-ossification; surface of the nasal rugoseexternally and marked by prominent spines internally; frontoparietals converge medially throughout theirlength, without exposing the fontanelle; anteriorly, the frontoparietal articulates with the nasal laterally andthe dermal sphenethmoid medially and anterolaterally, forming a slight supraorbital shelf; posterolaterallyfrontoparietal poorly developed and not articulate with the squamosal; posteriorly frontoparietal not form anoccipital crest covering the prootic; entire dorsal surface of the frontoparietal involved in integumentary cra-



nial co-ossification. Medially, the frontoparietal is rugose; irregular bony ridges radiate out from the centralrugosity toward the distal margins of the bone; dermal sphenethmoid triangular, centrally located at the ante-rior level of the orbit; sphenethmoid lies posteromedial to nasals and anteromedial to the frontoparietals; der-mal sphenethmoid is completely involved in integumentary-cranial co-ossification and its dorsal surfaceslightly rugose. Exoccipital and prootic completely fused, lacking a visible suture; anterodorsally exoccipitalspartially covered by frontoparietals; latter form the posterior border of the optic capsule; anterolaterally andlaterally, its crista parotica is in contact with the otic plate of the squamosal; dorsal and lateral surfaces of thesquamosal co-ossified. slightly rugose; zygomatic ramus robust and forms a part of the posterior margin of theorbit, it is connected to the maxillary and anterior rami of the pterygoid; optic ramus of squamosal poorlydeveloped, smaller than the zygomatic ramus and not extend posteriorly to the crista parotica; zygomaticramus articulates with the crista parotica, not with the frontoparietal; ventral arm of the squamosal moderatelyrobust; ventrally, it terminates between the quadratojugal laterally and the pterygoid medially, and dorsal tothe quadrate process. Pterygoid moderately robust; the anterior ramus lies adjacent to the posterior maxillaryprocess at a level just posterior to the orbitonasal foramen; medial ramus short, poorly developed and notarticulate with the prootic medially; posterior ramus robust and diverges medially from the maxillary to artic-ulate with the quadrate process and ventral arm of the squamosal posteriorly. Quadratojugal well developed;anterior end lies medially, adjacent to the maxillary at the level of the optic foramen; columella appearsbetween the levels of the prootic and anterior acoustic foramen.

FIGURE 3. Trachycephalus mambaiensis, MZUSP 135717 (paratype), adult male from the vicinity of Santa Edwiges Ihydroelectric power plant dam, Municipality of Mambaí, State of Goiás, Brazil. Note the small reddish spots disposed onthe dorsum and flank.



FIGURE 4. Trachycephalus mambaiensis, MZUSP 135715 (holotype), (A) hand in palmar view, (B) foot in plantarview. Scale = 10 mm.

Comparison with other species. The skull of T. mambaiensis is similar to T. nigromaculatus (Trueb,1970, plate 6a and 6b), differing from the latter by the less ossified conditions of the nasals, premaxillaries,and especially frontoparietals. The small development of the frontoparietals, failing to cover the prootics pos-teriorly, is a characteristic that easily differentiates the new species from the other species of the genus withossified skulls (Bokermann, 1966; Trueb, 1970). Trachycephalus mambaiensis can be further distinguishedfrom the the remaining species of Trachycephalus (all previously included in the genus Phrynohyas) by thepresence of a dermal sphenethmoid and by the less developed labial flange (see Trueb, 1970, Table 1). Theoverall characteristic of the skull of T. mambaiensis is intermediary between the condition shown by the spe-cies of Trachycephalus with an ossified skull and the ones without dermal ossifications (Fig. 6).

Distribution. The species is currently known only from the type-locality. However, it is possible that itsdistribution includes the upper rio Corrente region and its tributaries where several large fragments of naturalCerrado and gallery forests are still well preserved.

Etymology. A specific epithet in allusion to the municipality of Mambaí, where the species was collected.



FIGURE 5. Trachycephalus mambaiensis, MZUSP 135713 (paratype), (A) dorsal and (B) ventral views of the skull.

FIGURE 6. Dorsal view of skulls of Trachycephalus. (A) T. imitatrix (MZUSP 108928); (B) T. mambaiensis (MZUSP135713); (C) T. atlas (MZUSP 119815); (D) T. nigromaculatus (CHUNB 44221); (E) T. venulosus (MZUSP 138319);(F) T. coriaceus (MZUSP 60165); (G) T. mesophaeus (MZUSP 95705); (H) T. resinifictrix (MZUSP 69931).

Discussion

The northeastern State of Goiás is a typical Cerrado-Caatinga transition area, with intense fragmentation dueto agricultural practices (mainly soy bean and sugar cane plantations). The region has also suffered recently



from significant losses of natural habitats due to a drastic increase of interest for hydroelectric power produc-tion. As a consequence of hydroelectric activities, a number of rescue operations have been conducted onrequest of the Federal environmental agencies to monitor faunal diversity in the impacted natural habitats.During the flooding of the reservoir a faunal rescue operation is usually carried out to retrieve animals trappedwithin the flooded areas, resulting in the gathering of important vertebrate collections that offer a rare oppor-tunity to discover significant numbers of new taxa. Unfortunately, most of the details of habitat preferencesare lost during the process of flooding and inventory, which are generally tied to the planning of the construc-tion of the hydroelectric power plant. The new species described herein comes from a region that is alreadyunder intense environmental pressure, rendering urgent a continuous monitoring of the populations in order toascertain its conservation status.

The specialized vocal sac structure of Phrynohyas and Trachycephalus was viewed by Trueb andDuellman (1971) as an indication of their close affinities. Traditionally, the genus Trachycephalus was com-posed by species with highly ossified skulls and poorly developed parotoid glands, whereas species in thegenus Phrynohyas retained less ossified skulls and developed parotoid glands. Faivovich et al. (2005) haveshown that this division is artificial and the genus Phrynohyas should be synonymized with Trachycephalus,in order to render the group monophyletic. The intermediary morphology of the skull of T. mambaiensis, thatlies between the condition shown by the species of Trachycephalus with a well-ossified skull and the oneswithout dermal ossifications, corroborates the taxonomic decision made by Faivovich et al. (2005).

Acknowledgments

The authors are grateful to Guarino R. Colli (CHUNB) for allowing examination of specimens under his care,to José P. Pombal Jr. and an anonymous reviewer for their careful reviews of the manuscript, and to ItamarJunior Tonial for providing the photograph of the live specimen. We are also grateful to Mario de Pinna andEduardo G. Baena, who provided access and support to the Faxitron x-ray equipment. This research was sup-ported by grants from Fundação de Amparo à Pesquisa do Estado de São Paulo (BIOTA-FAPESP grant num-ber 02/13602–4) and from Conselho Nacional de Desenvolvimento Científico e Tecnológico to HZ (CNPqgrant number 303785/2004–7).

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Appendix. Specimens examined

Trachycephalus atlas (N=20)

BRAZIL: Bahia: Planalto Baiano: MZUSP 57099-57115; Maracás: MZUSP 119815-119818 (MZUSP 119815 skullexamined).

Trachycephalus coriaceus (N=8)

BRAZIL: Amazonas: Reserva INPA-WWF: MZUSP 60165 (skull examined); Pará: Serra de Kukoinhokren: MZUSP90900-90902; Rondônia: MZUSP 57345; Cachoeira de Nazaré: MZUSP 63810-63811; Mato Grosso: Apiacás:MZUSP 80811.

Trachycephalus imitatrix (N=20)

BRAZIL: São Paulo: São José do Barreiro: Campo de Fruticultura da Bocaina: MZUSP 108917-108936 (MZUSP108928 skull examined).

Trachycephalus jordani (N=10)

ECUADOR: Pichicha: Rio Palenque: MZUSP 56344-56353.

Trachycephalus lepidus (N=4)

BRAZIL: São Paulo: São Miguel Arcanjo: MZUSP 136548-136551.

Trachycephalus mesophaeus (N=10)

BRAZIL: Paraná: Caiobá: MZUSP 124477; Rio de Janeiro: Angra dos Reis: MZUSP 8995; Represa Rio Grande:MZUSP 95705-95712 (MZUSP 95705 skull examined).

Trachycephalus nigromaculatus (N=10)

BRAZIL: Espírito Santo: Presidente Kennedy: CHUNB 25037-25040; Goiás: São Domingos: CHUNB 21868, CHUNB35471, MZUSP 12625; Minas Gerais: Corinto: CHUNB 43212; Buritizeiro: CHUNB 43486, CHUNB 44221 (skullexamined).

Trachycephalus resinifictrix (N=14)

BRAZIL: Amazonas: Reserva INPA-WWF: MZUSP 57345; Reserva Ducke: MZUSP 84541, MZUSP 69929-69931(MZUSP 69931 – skull examined); Porto Alegre: MZUSP 60119-60122; Mato Grosso: Vila Rica: MZUSP 91907-91910; Pará: Serra de Kukoinhokren: MZUSP 71146.

Trachycephalus venulosus (N=11)

BRAZIL: Goiás: MZUSP 138319 (skull examined), MZUSP 138471-80.

A new species of Trachycephalus (Amphibia, Anura, Hylidae) from the State of Goiás, Brazil

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