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A New Species of Proceratophrys (Anura: Cycloramphidae) from MidwesternBrazilAuthor(s): Robson W. Ávila , André Pansonato , and Christine StrüssmannSource: Journal of Herpetology, 46(4):466-472. 2012.Published By: The Society for the Study of Amphibians and ReptilesDOI: http://dx.doi.org/10.1670/11-038URL: http://www.bioone.org/doi/full/10.1670/11-038

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Journal of Herpetology, Vol. 46, No. 4, 466–472, 2012Copyright 2012 Society for the Study of Amphibians and Reptiles

A New Species of Proceratophrys (Anura: Cycloramphidae) from Midwestern Brazil

ROBSON W. AVILA,1,2 ANDRE PANSONATO,3 AND CHRISTINE STRUSSMANN4

1Universidade Regional do Cariri, Centro de Ciencias Biologicas e da Saude, Departamento de Ciencias Biologicas, Campus do Pimenta, Rua Cel.Antonio Luiz, 1161, Bairro do Pimenta, CEP 63105-100, Crato, Ceara, Brazil

3Pos-Graduacao em Biologia Animal, Universidade Estadual Paulista–UNESP. R. Cristovao Colombo, 2265, Jardim Nazareth, CEP 15054-000,Sao Jose do Rio Preto, Sao Paulo, Brazil

4Departamento de Ciencias Basicas e Producao Animal, Faculdade de Agronomia, Medicina Veterinaria e Zootecnia,Universidade Federal de Mato Grosso (UFMT). Av. Fernando Correia da Costa, 2367, Boa Esperanca,

CEP 78060-900, Cuiaba, Mato Grosso, Brazil

ABSTRACT.—We describe adult morphology, advertisement call, and natural history aspects of a new frog species from riparian habitats

alongside intermittent headwater streams in southwestern Cerrado, the second largest biome in Brazil. Presently known from three localities in

Mato Grosso State, the new species belongs to the Proceratophrys cristiceps group. It is characterized mainly by small size (adult snout–vent

length approximately 46 mm), snout rounded in dorsal view and obtuse in lateral view, absence of flared lips, dorsal skin granular, and absenceof both postocular swellings and prominent palpebral appendages. The advertisement call of the new species consists of a single note with 19–

25 pulses. Calls are repeated at mean intervals of 0.7 sec. Mean dominant frequency is 1,250.2 Hz, with ascendant modulation.

The genus Proceratophrys Miranda-Ribeiro, 1920 presentlycomprises 25 species, distributed in Brazil, Argentina andParaguay (Cruz and Napoli, 2010; Avila et al., 2011; Martinsand Giaretta, 2011). Three species groups have been recognized:the Proceratophrys boiei group (sensu Prado and Pombal, 2008), theProceratophrys bigibbosa group (sensu Kwet and Faivovich, 2001),and the Proceratophrys cristiceps group (sensu Giaretta et al., 2000).Proceratophrys schirchi Miranda-Ribeiro, 1937 is not currentlyassociated to any of these groups, although Amaro et al. (2009)suggested its relationship with the group of P. cristiceps.

The P. cristiceps group is characterized by the absence ofboth postocular swellings and short palpebral appendages(sensu Giaretta et al., 2000). Seven species are currentlyrecognized in this group: two from Amazonian sites ofwestern Brazil (Proceratophrys concavitympanum Giaretta,Bernarde, and Kokubum 2000 and Proceratophrys strussmannaeAvila, Kawashita-Ribeiro, and Morais 2011), and the remain-ing five species distributed in the Cerrado domain of centraland southeastern Brazil (P. cristiceps Muller, 1883, Procera-tophrys goyana Miranda-Ribeiro, 1937, Proceratophrys cururuEterovick and Sazima, 1998, Proceratophrys moratoi Jim andCaramaschi, 1980, and Proceratophrys vielliardi Martins andGiaretta, 2011). Proceratophrys cristiceps is also found in theCaatinga from northeastern Brazil (Giaretta et al., 2000; Vieiraet al., 2008).

Boulenger (1903) examined the specimens of amphibians andreptiles collected by M. A. Robert during the Percy Sladenexpedition to central Brazil and incorrectly assigned specimens ofProceratophrys obtained at Chapada dos Guimaraes, Mato GrossoState, to P. cristiceps. During herpetological field studies conductedin this and in other two localities in Mid-Southern Mato Grosso,we obtained specimens of Proceratophrys that we recognize as apreviously undescribed taxon. Herein, we describe the newspecies that we assign to the P. cristiceps group.

MATERIALS AND METHODS

Specimens examined are deposited at the ‘‘Colecao Zoologicade Vertebrados da Universidade Federal de Mato Grosso,’’

Cuiaba, Brazil (UFMT), ‘‘Museu de Zoologia Adao Jose

Cardoso,’’ Universidade Estadual de Campinas, Campinas,

Brazil (ZUEC), and ‘‘Museu Nacional,’’ Rio de Janeiro, Brazil

(MNRJ). Specimens used for comparisons are listed in Appendix

1. For comparisons, data from congeneric species appears in

parentheses. Data for P. cururu appear in Eterovick and Sazima

(1998); for P. moratoi in Jim and Caramaschi (1980), and

Brasileiro et al. (2008), and for P. vielliardi are from Martins

and Giaretta (2011).

The following measurements (in millimeters) were taken with

digital calipers, to the nearest 0.01 mm, according to Heyer et al.

(1990) and Eterovick and Sazima (1998): ED (eye diameter);

END (eye to nostril distance); F (foot length), FL (forearm

length); H (hand length); HL (head length); HW (head width);

ID (internarial distance); IML (inner metatarsal tubercle length);

IND (internarial distance); SVL (snout–vent length); TD

(tympanum diameter); TF (tarsus–foot length); TH (thigh

length); TL (tibia length); UEW (upper eyelid width). Addition-

ally, the following characters were included: NT (number of

tubercles on the ventral surface of the left arm), NIT (number of

tubercles on inner surface of left arm), and NTP (number of

tubercles on upper eyelid). Webbing formula follows Savage

and Heyer (1997).

Advertisement calls of two adult males were recorded by

Christine Strussmann with a Sony TCM 5000-EV cassette tape

recorder, equipped with a Yoga EM-9600 external directional

microphone. Air temperature was measured using a standard

mercury thermometer of 0.18C. Tape recordings were digitized

at 44.1 kHz, resolution of 16 bits (FFT 1024), with Cool Edit 96

(eSyntrillium) and Raven Pro 1.3. They are archived as .wav

files in the ‘‘Banco de Registros Bioacusticos,’’ housed at

‘‘Laboratorio de Herpetologia do Instituto de Biociencias’’ of

the ‘‘Universidade Federal de Mato Grosso’’ (IB-UFMT, Cuiaba,

Mato Grosso, Brazil), under accession numbers LH 18A-01

(unvouchered) and LH 18A-02 (voucher specimen UFMT 1747).

Terminology for acoustic parameters follows Heyer et al. (1990)

and Martins and Jim (2003). The following seven call variables

were examined: note duration, internote intervals, mean rate of

notes per minute, number of pulses, pulse duration, interpulse

intervals, and dominant frequency.

2Corresponding Author. E-mail: robsonavila@gmail.comDOI: 10.1670/11-038

RESULTS

Proceratophrys huntingtoni sp. nov.

Orthochresonym: Proceratophrys cristiceps: Boulenger, 1903:69

Holotype.—UFMT 1745, an adult male collected on 29 August

2000 by Christine Strussmann during faunal monitoring activities

at Manso Hydroelectric Power Plant, Municipality of Chapada

dos Guimaraes, Mato Grosso State, central Brazil. The specimen

was calling from inside a grass tussock, near the headwaters of a

small stream (148520S; 558480W), after heavy rain.

Paratopotypes.—A total of seven paratopotypes: 2 adult females

(UFMT 1748 and 11134), and 5 adult males (UFMT 1746, 1747,

1749, 11133, and 11135) collected on 29 August 2000 by Christine

Strussmann.

Paratypes.—A total of 14 specimens: 5 adult females (UFMT

6698, 6699, 6700, 6702, and 6704) and 5 adult males (UFMT 6695–

6697, 6701, and 6703) collected on 2–3 September 2006 by Marcos

A. Carvalho at Fazenda Entre Rios (‘‘Jonas Pinheiro’’ small

Hydroelectric Power Plant), municipality of Santo Antonio do

Leverger, Mato Grosso State, Brazil; 3 adult males (UFMT 11129–

11131), and 1 adult female (UFMT 11132) collected on 27

November 2007 by Everaldo S. Moraes at Gruta da Cantina,

municipality of Nobres, Mato Grosso State, Brazil; one adult male

(UFMT 12635) collected on 20 July 2011 by Andre Pansonato and

FIG. 1. Holotype of Proceratophrys huntingtoni sp. nov., adult male (UFMT 1745): (A) Dorsal view; (B) ventral view; (C) ventral view of right hand;(D) ventral view of left foot; (E) lateral view of the head. Scale = 10 mm. Color reproduction supported by the Thomas Beauvais Fund.

NEW SPECIES OF PROCERATOPHRYS 467

Christine Strussmann at Vale do Uatuma, municipality ofChapada dos Guimaraes, Mato Grosso State, Brazil.

Diagnosis.—The new species is characterized by: small size(31.75–38.67 mm SVL in males [N = 14]; 33.04–45.69 mm SVL infemales [N = 8]), in comparison to congenerics; snout rounded indorsal view, obtuse in lateral view, dorsal skin granular, footlength representing approximately 40% of SVL, absence of bothpostocular swellings and prominent palpebral appendages.These last two characteristics lead us to consider the new speciesas a member of the Proceratophrys cristiceps group (sensu Giarettaet al., 2000).

Proceratophrys huntingtoni sp. nov. can be distinguished fromthe other species in the P. cristiceps group by the followingcharacters: from P. cristiceps, by smaller SVL (vs. 40.62–49.09mm in 5 males; 50.29–53.84 mm in 5 females), dorsal crests lessdefined beyond the sacral region (vs. more defined until thecoccyx), and snout obtuse in lateral view (vs. vertical); from P.concavitympanum, by smaller SVL (vs. 45.76–49.23 mm in 3males; 53.64–60.35 mm in 5 females), flared lips absent (vs. withflared lips), and inner metatarsal tubercle with dark brown color(vs. not colored); from P. cururu, by smaller SVL (vs. 36.5–43.1mm in males; 47.0–53.9 mm in females); from P. goyana, bysmaller SVL in males (vs. 43.41–46.3 mm; N = 2), snout obtusein lateral view (vs. vertical), dorsal crests interrupted at thesacral region (vs. more defined extending to the coccyx), andcanthal crests well defined (vs. not evident); from P. moratoi, byslightly larger SVL (vs. 24.7–30.9 mm in males; 32.1–41.6 mm in

females), snout obtuse in lateral view (vs. vertical), and longerlegs (vs. mean TH 10.9 mm, range 10.2–11.6 mm; mean TL 9.9,range 9–10.7 mm); from P. strussmannae, by smaller SVL (vs.41.1–47.3 mm in males; 52.7–59.8 mm in females); from P.vielliardi, by smaller SVL in males (vs. 39.1–41.9 mm in males),and presence of dorsal crests interrupted at the sacral region (vs.absent).

Description of the Holotype.—Robust and stout body, head widerthan long, head width representing 45.6% of SVL, snout roundedin dorsal view and obtuse in profile (Fig. 1A,E), eye diameter 20%larger than distance of eye to nostril, slightly smaller than uppereyelid length. Nostrils elliptical; canthal crests present on thecanthus rostralis; 7 warts of equal size located at the margin ofupper eyelid; a cutaneous dorsal crest from middle upper eyelidto sacral region, not reaching the coccyx; a row of three enlargedwarts, initiating slightly above the corner of mouth, with the lastat the level of mouth; a row of 8 tubercles on the lower jaw. Tworows of parallel warts (formed by 7 and 4 warts) extending fromthe end of hand to two-thirds of the length of the forearm.Fingers free, distally unexpanded, relative length 4 < 2 < 1 < 3;inner carpal tubercle oval in shape; outer carpal tubercle oblique,divided; subarticular and supernumerary finger tubercles en-larged, rounded (Fig. 1C). Toes with unexpanded tips, relativelength 1 < 2 < 5 < 3 < 4; inner metatarsal tubercle piriform, witha dark brown color in the edge; outer metatarsal tuberclerounded, very small (Fig. 1D); a ridge of three warts in the tarsus,from beyond to the outer metatarsal tubercle; toes webbed

TABLE 1. Measurements (in millimeters) of male (including holotype) and female type specimens of Proceratophrys huntingtoni sp. nov. Means are6 1 standard deviation, ranges are in parentheses. Character abbreviations are defined in the Material and Methods section.

Character Male (N = 14) Female (N = 8)

SVL 34.68 6 2.07 (31.75–38.67) 41.06 6 5.16 (33.04–45.69)HL 13.87 6 1.10 (12.41–15.95) 16.51 6 1.95 (13.19–18.70)HW 15.84. 6 0.88 (14.25–16.97) 18.34 6 2.56 (15.08–21.74)ED 4.33 6 0.47 (3.85–5.21) 4.83 6 0.42 (4.26–5.45)END 3.08 6 0.34 (2.53–3.54) 3.94 6 0.74 (2.64–4.66)TD 2.01 6 0.34 (1.53–2.68) 2.15 6 0.40 (1.67–2.71)TH 13.46 6 1.18 (11.24–15.87) 14.84 6 1.68 (12.47–17.01)TF 18.03 6 1.23 (16.37–20.10) 21.64 6 3.11 (17.50–25.77)F 14.36 6 0.97 (12.83–15.86) 16.88 6 2.45 (13.89–19.90)H 9.31 6 0.56 (8.31–10.30) 10.91 6 1.56 (8.74–12.90)IND 2.41 6 0.30 (1.74–3.17) 2.99 6 0.82 (1.90–3.85)TL 13.36 6 0.86 (12.06–14.84) 15.18 6 1.48 (12.93–16.77)FL 8.16 6 0.42 (7.45–8.73) 9.12 6 0.84 (8.18–10.13)IML 2.76 6 0.31 (2.25–3.24) 3.37 6 0.50 (2.54–4.06)UEW 4.10 6 0.30 (3.61–4.86) 4.63 6 0.56 (4.12–5.49)

FIG. 2. Dorsal (A) and ventral (B) views of a live specimen (UFMT 12635) of Proceratophrys huntingtoni sp. nov. collected on 20 July 2011 at Vale doUatuma, Chapada dos Guimaraes, Mato Grosso, Brazil. Color reproduction supported by the Thomas Beauvais Fund.

468 R. W. AVILA ET AL.

basally, webbing formula: I 1-–2+ II 2+–3 III 3–2 ½ IV 3+–2 V.

Foot length equals 23% of SVL. Dorsal skin basically granular,

with scattered discrete warts; chest and belly covered with

granulated skin. Dentigerous process of vomers oblique and

barely separated, between choanae; vocal slits present, lateral to

tongue. Tongue ovoid, covering the entire floor of mouth,

notched posteriorly.

Measurements of the Holotype (in Millimeters).—SVL 33.90; HL

13.46; HW 15.47; ED 3.70; END 2.96; TD 2.15; TH 13.55; TF 18.72;

F 14.02; H 9.49; IND 2.45; TL 14.05; FL 8.25; IML 2.64; UEW 4.09.

Coloration.—In life, five dark brown bands are present on each

side of the face, each one separated by grayish-brown stripes. A

dark brown stripe on canthus rostralis. There are 2–3 dark brown

transverse stripes both on forearms, thighs and tibias. Dorsum

predominantly light brown, with darker irregular patches in

dorsolateral region and a clearer ‘‘X’’ on the back, from the

supraocular region to the rump. Two triangular dark spots on

interocular region. Irregular black reticulations all over the iris:

over the golden middle section of its dorsal half, over the whitish

middle section of its lower half, and over chestnut brown of

anterior and posteriormost sections of the iris. Ventral color

pattern not recorded in life.

Variation.—Sexual dimorphism is apparent, with femalesattaining greater SVL (Table 1). Adult males have a blackpigmented vocal sac (Fig. 1B). Dorsal coloration is highly variableand there is a variation also in vermiculation pattern on belly.Four specimens (UFMT 6698, 6700, 6703, 6704) out of 22 have alight brown belly, without vermiculations.

The coloration of a recently collected male specimen (UFMT12635) was noted as follows: belly light vermiculations, chestand gular vermiculations, an evident patch in the innermostportion of the anterior ventral surface of the thighs. Littlechange in coloration is noted in relation to preserved specimens,with the exception of some apparent darkening of the brownmarkings (Fig. 2).

Advertisement Call.—The advertisement call of P. huntingtoni isdescribed based on 2 males recorded on 15 December 2000,respectively at 1815 h (unvouchered specimen) and 1830 h(UFMT 1747), at air temperature of 248C. Both calling males werefound semiburied, inside a grass tussock, in a narrow galleryforest, at the margin of a small, intermittent oligotrophicheadwater stream (14855026 00S; 55848020 00W) amid a matrix ofopen Cerrado vegetation (see below).

The advertisement call of P. huntingtoni (Fig. 3) consists of asingle note with 19–25 pulses (N = 103), and ascendant

FIG. 3. Advertisement call of Proceratophrys huntingtoni sp. nov. (UFMT 1747), recorded in the municipality of Chapada dos Guimaraes, MatoGrosso State, on 15 December 2000, 1830 h, air temperature 248C. (A) Oscillogram; (B) audio spectrogram.

TABLE 2. Measurements of acoustic parameters for two males of Proceratophrys huntingtoni sp. nov. Values are: mean 6 1 standard deviation, andrange, in parentheses. LH = Banco de Registros Bioacusticos, Laboratorio de Herpetologia, IB-UFMT. LH 18A-01 (unvouchered) and LH 18A-02(voucher specimen UFMT 1747). N = number of calls analyzed.

LH 18A–01 (N = 46) LH 18A–02 (N = 57) Mean (N = 103)

Note duration (sec) 0.275 6 0.017 (0.2–0.32) 0.271 6 0.01 (0.24–0.3) 0.27 6 0.01 (0.2–0.3)Internotes interval (sec) 0.7 6 0.38 (0.37–2) 0.7 6 0.35 (0.34–2.38) 0.7 6 0.36 (0.3–2.3)Dominant frequency (Hz) 1284.9 6 23.6 (1263.8–1344.5) 1222.1 6 47.8 (1095–1316.7) 1250.2 6 49.9 (1095–1344.5)Number of pulses 22 6 1.3 (20–25) 20.8 6 0.8 (19–23) 21.3 6 1.2 (19–25)Pulse duration 0.007 6 0.001 (0.006–0.009) 0.007 6 0.001 (0.005–0.009) 0.007 6 0.001 (0.005–0.009)Interpulses interval (sec) 0.005 (0.004–0.006) 0.0045 (0.004–0.006) 0.0046 6 0.0006 (0.004–0.006)Mean rate of notes/min 64.25 6 11.9 65.15 6 6.26 64.63 6 9.17

NEW SPECIES OF PROCERATOPHRYS 469

frequency modulation. Calls have a mean duration of 0.27 6

0.01 (range 0.2–0.3 sec) and are repeated at mean intervals of 0.7sec. Notes were emitted at a rate of 64.6/min. Pulses have meanduration of 0.007 sec (except for the last pulse, which variesfrom 0.16–0.2 sec) and mean intervals of 0.005 sec. The meandominant frequency was 1,250.2 Hz (Table 2).

Natural History.—The new species inhabits narrow galleryforests alongside intermittent headwater streams amidst Cerrado.Males were heard calling near the margins of such intermittentstreams both in relatively plain terrains and on the slopes ofgranitic (Santo Antonio do Leverger), limestone (Nobres), andsandstone formations (Chapada dos Guimaraes). Males wererecorded in a small tributary of ‘‘Corrego Bananal,’’ which flowsinto the Casca River (which belongs to Manso River sub basin,and ultimately to the Cuiaba River Basin and Upper ParaguayRiver Basin). The small stream is situated at Fazenda Morro doChapeu, Chapada dos Guimaraes municipality, Mato GrossoState.

Proceratophrys huntingtoni is a nocturnal, terrestrial species.The climate in the geographic range of the new species isgenerally hot and semihumid, with two well-marked seasons:the rainy season, from October to April, and the dry season,from May to September. Mean temperature in Chapada dosGuimaraes region is 268C, and mean monthly relative airhumidity varies from 60–83%, with a drop to approximately30% in July and August (Alho, 2000). Total annual precipitationis 1,800 mm, on average (Brasil, 1992).

Distribution and Conservation Concerns.—Proceratophrys hunting-toni is known from three municipalities in Mid-Southern MatoGrosso State, all of them in the Cerrado domain (Fig. 4). TheCerrado encompasses approximately 2 million km2, being thesecond largest vegetational domain in Brazil (Ratter et al., 1997).Despite their biological importance (Myers et al., 2000), naturalareas of Cerrado are being greatly disturbed by the advance ofthe booming agricultural frontier, with more than 2 million ha ofpristine habitats destroyed each year (Klink and Machado, 2005).Additionally, many huge areas of Brazilian Cerrado have beenlost because of the establishment of hydroelectric impoundments(e.g., Manso powerplant, 427 km2, Mato Grosso State; Serra daMesa, 1,780 km2, Goias State; see Alho, 2000, and Brandao and

Araujo, 2008, respectively). Also, complexes of townhouses andother facilities are usually built near the margins of newreservoirs, as is the case at Manso Power Plant, increasing landdivision and habitat losses.

Etymology.—The specific epithet honors Herbert HuntingtonSmith (1851–1919), an American naturalist and amateur conchol-ogist, who signed himself ‘‘H. Huntington’’ in some of hischronicles. Smith traveled into southern Mato Grosso andcollected zoological specimens in the small village of Chapadados Guimaraes between 1882 and 1885. Specimens of amphibiansand reptiles obtained by Smith in Chapada were sent to theherpetologist Edward Drinker Cope, from the Academy ofNatural Sciences (Philadelphia, PA), who described many newtaxa based in that material (Cope, 1887). Interesting informationon geology, landscapes, and biogeography of southern MatoGrosso were published in the book ‘‘Do Rio de Janeiro a Cuyaba:notas de um naturalista’’ (Smith, 1922). Smith’s writings from theperiod spent in Chapada, however, were seemingly neverpublished, as was his original intention.

DISCUSSION

Proceratophrys huntingtoni is the sixth species of the P. cristicepsgroup recorded for the Cerrado biome. In addition, at least onespecies in the same group, Proceratophrys concavitympanum, wasoriginally described from Amazonian sites and lately recordedin forested habitats in central Cerrado (Pavan, 2007).

In the remaining two species groups within the genusProceratophrys, species in the P. boiei group (12 species in total)are mostly restricted to the Atlantic Forest, except for P.rondonae, described from Amazonian sites of the Guapore Riverdrainage, in the Brazilian state of Rondonia (Prado and Pombal,2008). Contrarily, in the P. bigibbosa species group (comprisingfour species), most species are restricted to subtropical regionsof southern Brazil and Argentina (Kwet and Faivovich, 2001).

Note duration of the advertisement call of P. huntingtoni issimilar to that in P. moratoi and in P. concavitympanum (Brasileiroet al., 2008; Santana et al., 2010, respectively). However, it isstrikingly different from note duration in other species ofProceratophrys also belonging to the P. cristiceps group (e.g., P.cristiceps, P. cururu and P. vielliardi; Table 3). It is very distinct,also, from note duration in species of the P. bigibbosa group: P.avelinoi (Kwet and Baldo, 2003), P. brauni (Kwet and Faivovich,2001), and also in P. paviotti (Cruz et al., 2005) and P. sanctaritae(Cruz and Napoli, 2010), species in the P. boiei group.

The dominant frequency of the advertisement call of P.huntingtoni is similar to that in P. moratoi (Brasileiro et al., 2008)and P. vielliardi (Martins and Giaretta, 2011), being lower thandominant frequency in P. avelinoi (Kwet and Baldo, 2003), andconsiderably higher than dominant frequency recorded for P.cristiceps (Nunes and Junca, 2006), P. concavitympanum (Santanaet al., 2010), P. cururu (Eterovick and Sazima, 1998), P.melanopogon (Mangia et al., 2010), P. sanctaritae (Cruz andNapoli, 2010), and P. paviotti (Cruz et al., 2005).

The number of pulses in the call of P. huntingtoni is lower thanin P. cristiceps (Nunes and Junca, 2006), P. cururu (Eterovick andSazima, 1998), and P. sanctaritae (Cruz and Napoli, 2010) andhigher than in P. vielliardi (Martins and Giaretta, 2011). Noterepetition rate is higher in the new species than in P. moratoi(Brasileiro et al., 2008) but lower than in P. vielliardi (Martins andGiaretta, 2011; Table 3). The ascendant frequency modulationfound in the advertisement call of P. huntingtoni is known tooccur in two other species: P. cururu, and P. paviotti (Eterovick

FIG. 4. Known geographic distribution of Proceratophrys huntingtonisp. nov. Circles: (1) Nobres municipality; (2) Santo Antonio do Levergermunicipality. The star represents the type locality: Chapada dosGuimaraes, Mato Grosso State, Brazil.

470 R. W. AVILA ET AL.

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NEW SPECIES OF PROCERATOPHRYS 471

and Sazima, 1998; Cruz et al., 2005), being absent in P. cristiceps(Nunes and Junca, 2006) (Table 3).

With the addition of P. huntingtoni, the number of species inthe P. cristiceps group raises to eight. Because of deficientinformation in the original description of many species in thisgroup, a thorough revision of it, as previously made for the P.biggibosa and P. boiei groups by Prado and Pombal (2008), andby Kwet and Faivovich (2001), respectively, is badly needed.Proceratophrys goyana, for example, was described based only ontwo specimens from the state of Goias, and erroneouslyillustrated with a figure of P. cristiceps (see Miranda-Ribeiro,1937).

Acknowledgments.—Thanks are due to Marcos A. Carvalho,Jose P. Pombal Jr., and Luis F. Toledo for providing access to theColecao Zoologica de Vertebrados da Universidade Federal deMato Grosso, Museu Nacional do Rio de Janeiro, and Museu deZoologia Adao Jose Cardoso, of the Universidade Estadual deCampinas, respectively. Two anonymous reviewers and ErinMuths made valuable comments and suggestions to improvethe quality of the paper.

LITERATURE CITED

ALHO, C. J. R. 2000. Paisagens e enfoque. In: C. J. R. Alho, (Coord.),Fauna Silvestre da regiao do rio Manso, MT. pp. 19–29. Ministerio doMeio Ambiente, Edicoes IBAMA and Centrais Eletricas do Norte doBrasil, Brasılia.

AMARO, R. C., D. PAVAN, AND M. T. RODRIGUES. 2009. On the genericidentity of Odontophrynus moratoi Jim and Caramaschi, 1980 (Anura,Cycloramphidae). Zootaxa 2071:61–68.

AVILA, R. W., R. A. KAWASHITA-RIBEIRO, AND D. H. MORAIS. 2011. A newspecies of Proceratophrys (Anura: Cycloramphidae) from westernBrazil. Zootaxa 2890:20–28.

BOULENGER, G. A. 1903. List of the batrachians and reptiles collected byM. A. Robert at Chapada, Matto Grosso, and presented by Mrs.Percy Sladen to the British Museum (Percy Sladen Expedition toCentral Brazil). Proceedings of the Zoological Society of London 2:69–70.

BRANDAO, R. A., AND A. F. B. ARAUJO. 2008. Changes in anuran speciesrichness and abundance resulting from hydroelectric dam floodingin Central Brazil. Biotropica 40:263–266.

BRASIL. 1992. Ministerio da Agricultura e Reforma Agraria. SecretariaNacional de Irrigacao. Departamento Nacional de Meteorologia.Normais Climatologicas (1961–1990). Brasılia.

BRASILEIRO, C. A., I. A. MARTINS, AND J. JIM. 2008. Amphibia, Anura,Cycloramphidae, Odontophrynus moratoi: distribution extension andadvertisement call. Checklist 4:382–385.

COPE, E. D. 1887. Synopsis of the Batrachia and Reptilia obtained by H.H. Smith in the Province of Mato Grosso, Brazil. Proceedings of theAmerican Philosophical Society 24:44–60.

CRUZ, C. A. G., AND M. F. NAPOLI. 2010. A new species of smooth hornedfrog, genus Proceratophrys Miranda-Ribeiro (Amphibia: Anura:Cycloramphidae), from the Atlantic Rainforest of eastern Bahia,Brazil. Zootaxa 2660:57–67.

CRUZ, C. A. G., G. M. PRADO, AND E. IZECKSON. 2005. Nova especie deProceratophrys Miranda-Ribeiro, 1920 do sudeste do Brasil (Amphib-ia, Anura, Leptodactylidae). Arquivos do Museu Nacional 63:289–295.

ETEROVICK, P. C., AND I. SAZIMA. 1998. New species of Proceratophrys(Anura: Leptodactylidae) from southeastern Brazil. Copeia 1998:159–164.

GIARETTA, A. A., P. S. BERNARDE, AND M. N. C. KOKUBUM. 2000. A newspecies of Proceratophrys (Anura: Leptodactylidae) from the AmazonRain Forest. Journal of Herpetology 34:173–178.

HEYER, W. R., A. S. RAND, C. A. G. CRUZ, O. L. PEIXOTO, AND C. E. NELSON.1990. Frogs of Boraceia. Arquivos de Zoologia 31:231–410.

JIM, J., AND U. CARAMASCHI. 1980. Uma nova especie de Odontophrynus daregiao de Botucatu, Sao Paulo, Brasil (Amphibia, Anura). RevistaBrasileira de Biologia 40:357–360.

KLINK, C. A., AND R. B. MACHADO. 2005. Conservation of the BrazilianCerrado. Conservation Biology 19:707–713.

KWET, A., AND D. BALDO. 2003. Advertisement call of the leptodactylidfrog Proceratophrys avelinoi. Amphibia-Reptilia 24:104–107.

KWET, A., AND J. FAIVOVICH. 2001. Proceratophrys bigibbosa species group(Anura: Leptodactylidae), with description of a new species. Copeia2001:203–215.

MANGIA, S., D. J. SANTANA, AND R. N. FEIO. 2010. Advertisement call of thecycloramphid toad Proceratophrys melanopogon (Miranda-Ribeiro,1926). South American Journal of Herpetology 5:127–131.

MARTINS, L. B., AND A. A. GIARETTA. 2011. A new species of Proceratophrys(Amphibia: Anura: Cycloramphidae) from central Brazil. Zootaxa2880:41–50.

MARTINS, I. A., AND J. JIM. 2003. Bioacoustic analysis of advertisement callin Hyla nana and Hyla sanborni in Botucatu, Sao Paulo, Brazil.Brazilian Journal of Biology 63:507–516.

MIRANDA-RIBEIRO, A. 1937. Especies novas do genero ‘‘Stombus’’ da seriede appendices oculares reduzidos. O Campo 8(88):24.

MYERS, N., R. A. MITTERMIER, C. G. MITTERMIER, G. A. B. FONSECA, AND J.KENT. 2000. Biodiversity hotspots for conservation priorities. Nature403:853–858.

NUNES, I., AND F. A. JUNCA. 2006. Advertisement calls of threeleptodactylid frogs in the state of Bahia, northeastern Brazil(Amphibia, Anura, Leptodactylidae), with considerations on theirtaxonomic status. Arquivos do Museu Nacional 64:151–157.

PAVAN, D. 2007. Assembleias de repteis e anfıbios do Cerrado ao longoda bacia do rio Tocantins e o impacto do aproveitamento hidreletricoda regiao na sua conservacao. Unpubl. Ph.D. thesis. Instituto deBiociencias. Universidade. de Sao Paulo, Sao Paulo, Brazil.

PRADO, G. M., AND J. P. POMBAL JR. 2008. Especies de ProceratophrysMiranda-Ribeiro, 1920 com apendices palpebrais (Anura; Cyclo-ramphidae). Arquivos de Zoologia 39:1–85.

RATTER, J. A., J. F. RIBEIRO, AND S. BRIDGEWATER. 1997. The BrazilianCerrado vegetation and threats to its biodiversity. Annals of Botany80:223–230.

SANTANA, D. J., V. A. SAO-PEDRO, P. S. BERNARDE, AND R. N. FEIO. 2010.Descricao do canto de anuncio e dimorfismo sexual em Procera-tophrys concavitympanum Giaretta, Bernarde and Kokubum, 2000.Papeis Avulsos de Zoologia 50:167–174.

SAVAGE, J. M., AND W. R. HEYER. 1997. Digital webbing formulae foranurans: a refinement. Herpetological Review 28:131.

SMITH, H. H. 1922. Do Rio de Janeiro a Cuyaba: Notas de um naturalista.Melhoramentos, Sao Paulo, Brazil.

VIEIRA, K. S., C. ARZABE, M. I. M. HERNANDEZ, AND W. L. S. VIEIRA. 2008.An examination of morphometric variations in a Neotropical toadpopulation (Proceratophrys cristiceps, Amphibia, Anura, Cycloram-phidae). PLoS ONE 3(12):e3934.

Accepted: 26 October 2011.

APPENDIX 1

Specimens Examined

Proceratophrys concavitympanum.—UFMT 6808 (Colniza, Mato Gros-so); UFMT 6996 and 7825 (Juına, Mato Grosso); UFMT 7906 (Paranaıta,Mato Grosso); ZUEC 11387, 11388 (Espigao D’Oeste, Rondonia–holotype and paratype); ZUEC 14874, 14876, 16015 (Paranaıta, MatoGrosso), ZUEC 14950 (Jacareacanga, Para).

Proceratophrys cristiceps.—MNRJ 16472–16481.Proceratophrys bigibbosa.—ZUEC 10405–10406 (Sao Francisco de Paula,

Rio Grande do Sul).Proceratophrys goyana.—ZUEC 4110 (Picos, Piauı); ZUEC 4125

(Santana do Cariri, Ceara).Proceratophrys moratoi.—ZUEC 7031–7033 (Rubiao Jr., Botucatu, Sao

Paulo).Proceratophrys strussmannae.—UFMT 5859, 6659, 7869, 7872, 7874,

7876, 7878, 7880, 7882, 7885, 7886, 8319, 8320, 8377–8380 (Jauru and Valede Sao Domingos, Mato Grosso).

472 R. W. AVILA ET AL.