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New Species of Melanophryniscus (Anura: Bufonidae) from RioGrande do Sul, Southern BrazilAuthor(s): Marcos Di-Bernardo, Raúl Maneyro, Hamilton GrilloSource: Journal of Herpetology, 40(2):261-266. 2006.Published By: The Society for the Study of Amphibians and ReptilesDOI: http://dx.doi.org/10.1670/05-008.1URL: http://www.bioone.org/doi/full/10.1670/05-008.1

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New Species of Melanophryniscus (Anura: Bufonidae) fromRio Grande do Sul, Southern Brazil

MARCOS DI-BERNARDO,1 RAUL MANEYRO,1,2,3AND HAMILTON GRILLO

4

1Laboratorio de Herpetologia, Museu de Ciencias e Tecnologia and Faculdade de Biociencias,Pontifıcia Universidade Catolica do Rio Grande do Sul, Avenida Ipiranga 6681,

90619-900 Porto Alegre, Rio Grande do Sul, Brazil4Museu de Ciencias Naturais, Universidade do Vale do Taquari, Rua Avelino Tallini,

171, 95900-000 Lajeado, Rio Grande do Sul, Brazil

ABSTRACT.—A new bufonid toad is described. The species belongs to the genus Melanophryniscus, and may

be the largest species in this genus. Its size and external features (pale green in dorsal surface) resemble thoseof Melanophryniscus cambaraensis and Melanophryniscus macrogranulosus, from which it differs by the

absence of frontal swelling. At present, the new species is endemic to the southern border of the Brazilian

Southern Plateau, in central Rio Grande do Sul state.

The South American genus Melanophryniscuswas created by Gallardo (1961), based onMelanophryniscus stelzneri (type-species), Melano-phryniscus tumifrons, and Melanophryniscus mor-eirae. To date, this genus consists of 19 species(Kwet et al., 2005), which are distributed fromBolivia through Paraguay, north to Argentina,and south and southeast of Brazil to Uruguay(Frost, 2004).

Among the bufonids, this genus is consideredmonophyletic and basal (Graybeal and Can-natella, 1995; Haas, 2003). Three (Cruz and Car-amaschi, 2003) or four (Larson et al., 2003)phyletic groups have been defined based onexternal morphology and coloration. However,only the M. tumifrons group has a proposedsynapomorphy, the frontal swelling (Baldo andBasso, 2004).

The genus Melanophryniscus is widespreadin the state of Rio Grande do Sul (Brazil), whereit is represented by eight species included inthe M. stelzneri and M. tumifrons groups. Amongthem, Melanophryniscus cambaraensis and Mela-nophryniscus macrogranulosus (both in the M.tumifrons group) are endemic to the state. Aninth species, Melanophryniscus devincenzii, wascited for this state (Di-Bernardo et al., 2004)based on previous work (Cespedez and Motte,2001), but the presence of this species requiresconfirmation (no specimens are deposited inscientific collections). Herein we describe anew species of Melanophryniscus endemic to

Rio Grande do Sul, collected on the slopes ofthe southern border of the Brazilian SouthernPlateau.

MATERIALS AND METHODS

This description is based on 31 specimensfrom three series, deposited in the herpetologicalcollection of the Museu de Ciencias e Tecnologiaof the Pontifıcia Universidade Catolica do RioGrande do Sul (MCP), Brazil; Museu de CienciasNaturais da Fundacao Zoobotanica do RioGrande do Sul (MCN), Brazil; and Facultad deCiencias de la Universidad de la Republica(ZVCB), Uruguay. The frogs belonging to thefirst series were donated to the authors about2 months after collection. Six of them weremaintained in captivity by the collectors, eachdied on different dates (not recorded) and werefixed and stored in 10% formalin. Two speci-mens, an amplectant pair at the time of collecting,were donated alive. The second and third serieswere collected by the authors and collaborators.The following measurements (as in Baldo andBasso, 2004) were taken using vernier calipers tothe nearest 0.05 mm: snout–vent length (SVL),head length (HL), mandibular width (MW), eyediameter (ED), interorbital distance (IOD), snout-eye distance (SED), internarial distance (ID),upper forearm length (UF), lower forearm length(LF), thigh length (TL), leg length (LG), and footlength (FT). For each measure (N 5 31), mean,standard deviation (SD), and range are given(Table 1).

Illustrations of the head, hand, and footwere made using a stereoscopic microscope(Zeiss Stemi SV-11) and an attached cameralucida.

2 Present Address: Seccion Zoologıa Vertebrados,Facultad de Ciencias, Universidad de la Republica,Igua 4225, 11400 Montevideo, Uruguay; E-mail:rmaneyro@fcien.edu.uy

3 Corresponding Author.

Journal of Herpetology, Vol. 40, No. 2, pp. 261–266, 2006Copyright 2006 Society for the Study of Amphibians and Reptiles

Melanophryniscus admirabilis, sp. nov.Figure 1

Holotype.—MCP 7923 (Figs. 2–4), adult female,collected in Perau de Janeiro (528189W, 288519S),Municipality of Arvorezinha, Rio Grande do Sul,Brazil, by A. E. Oliveira on 8 January 2004.

Paratypes.—All adults and from the same lo-cality as the holotype: MCP 7922, male, and MCP7956-61 collected by A. E. Oliveira on 8 January2004; MCP 8062 (Fig. 1) and 8063 collected byC. Schwingel, L. B. de Oliveira, D. Goncalves, andH. Grillo on 19 September 2004; MCP 8068–70,8072, 8075–77, 8088–91, 8093–96, ZVCB 11531–33, MCN 13821–23 collected by M. Di-Bernardo,C. Di-Bernardo, C. D. N. Castro, G. F. Pontes,R. B. Oliveira, R. L. Balestrin, L. H. Cappellari,J. Frota, C. Zank, W. Vaz, C. Schwingel, L. B. deOliveira, D. Goncalves, and H. Grillo on 2 October2004.

Diagnosis.—M. admirabilis is the largest speciesbelonging to this genus (SVL range 5 40.28–29.48 mm). It is characterized by absence offrontal swelling, and presence of very well-developed glands in dorsal, lateral, and ventralsurfaces. The skin glands are very conspicuousand easily distinguishable because of their con-trasting coloration.

The new species can be differentiated from allother taxa in the M. stelzneri and M. moreiraegroups by its large size (adults usually reachmore than 30 mm in SVL), large size of the skinglands, presence of large glands on the lateraland ventral surfaces, and pale green dorsal col-oration (all characters are absent in the twogroups above). M. admirabilis differs from speciesin the M. tumifrons species group by the absenceof the frontal swelling. However, colorationresembles that of M. cambaraensis, from which itcan be distinguished by the contrasting colora-tion of the dorsal skin glands (yellowish in thenew species and green in M. cambaraensis). Thesize of the dorsal glands resembles another taxonin the M. tumifrons species group, M. macro-granulosus, but in this taxon, the dorsum ishomogeneously dark green (Escobar et al., 2004).

Description of the Holotype.—Body stout; headlonger than wide (Fig. 2); glands of the skindiminish in size in the dorsal surface of the head(absent from the interorbital region to the tip ofthe snout); neck indistinguishable from head;

snout truncated in dorsal and lateral views; nos-trils oval, located subterminally respect to thesnout and open in a lateral direction; canthusrostralis rounded and not evident; loreal regionconcave; eyes large; eyelid length less thaneyelid-nostril distance and than interorbital dis-tance; pupils rounded and black; tympanic mem-branes not visible; parotoid glands absent;external border of the upper maxillae wider thanthe external border of the mandibles (this lastencompassed by the former, that resembles anupper lip); upper lip with a medial and verysmall reentrant; maxillae and premaxillae eden-tate; vocal sac median subgular (externally blackwith few little yellowish glands); tongue narrowand elongate, with its posterior margin entire androunded; dorsal skin with very conspicuousglands from the neck to the vent (also conspic-uous, but smaller on lateral and ventral surfaces);fore and hind limbs slender with dorsal surfacescovered by the typical granulation describedabove; external metacarpal tubercle also con-spicuous (more than 2 mm in diameter); innermetacarpal tubercle smaller; fingers thick androunded at the tips; predominant hand formula:I , IV , II , III, (fingers I, II, and IV are sub-equal); hands without webbing but possessubarticular and palmar tubercles (Fig. 3); hindfeet with two metatarsal tubercles (inner severaltimes larger than the outer); feet with a poorlydeveloped interdigital membrane (particularlybetween fingers I, II, and III); foot formula: I ,II , V , III , IV; tips of the toes rounded;subarticular feet tubercles more conspicuousthan those on the forelimb digits (Fig. 4).

Measurements of the Holotype (in mm).—SVL 538.36, HL 5 10.81, MW 5 10.01, ED 5 4.04,IOD 5 6.20, SED 5 4.39, ID 5 3.31, UF 5 10.91,LF 5 8.88, TL 5 14.96, LG 5 13.27, FT 5 21.60.

Color in Life.—Predominant color of dorsalsurface green, which becomes paler, yellowish, inthe glands; chin and gular region black withsome yellow glands; chest and belly black, withirregular pale green or yellowish (sometimes red-centered spots) glands; ventral surface of thighsred; flanks darker than dorsum, with large yel-low rounded glands; fore and hind limbs green indorsal view; palms and soles bright red; pupillimited with a brown ring; iris gold and greenwith brown spots (Fig. 1).

TABLE 1. Summary statistics of the measures of the paratypes of Melanophryniscus admirabilis. Values aregiven in millimeters (N 5 30). See abbreviations in text.

SVL HL MW ED IOD SED ID UF LF TL LG FT

Mean 35.53 10.56 10.40 3.95 6.05 4.78 3.14 11.65 8.54 13.91 13.04 20.83SD 2.25 0.59 0.58 0.25 0.42 0.46 0.20 0.80 0.61 0.81 0.72 1.05Maximum 40.28 11.58 11.83 4.53 7.09 5.50 3.48 13.20 9.75 16.05 14.62 23.56Minimum 29.48 8.98 9.03 3.41 5.37 4.02 2.69 10.04 7.50 12.28 10.92 18.41

262 M. DI-BERNARDO ET AL.

FIG. 1. Melanophryniscus admirabilis sp. nov. (MCP 8062). (top) Lateral view; (bottom) ventral view. Scale bars5 5.0 mm.

NEW BRAZILIAN MELANOPHRYNISCUS 263

Color in Preservative.—Primary dorsal colora-tion grey and primary ventral coloration black;glands paler, mainly on the ventral surface,where they contrast deeply with the dark groundcoloration; inner surface of the thighs, palmsand soles yellowish.

Etymology.—The specific epithet refers to theadjective ‘‘admirable,’’ because of the beautifulcoloration of this toad.

Distribution.—M. admirabilis presently isknown only from the type locality in Rio Grandedo Sul, Brazil (Fig. 5).

Variation.—The dorsal and ventral color pat-terns are quite conservative, given that the spatialdistribution of the glands is different in eachspecimen. The extent and brightness of the redcoloration on the ventral surfaces of the thighs is

variable, being less evident in some specimens(e.g., MCP 7961). The amount of webbing on thefeet is variable, being absent in some specimens(e.g., MCP 7956). Compared to the holotype, thesnout is less truncated in three specimens (MCP7958, 7960, and 7961). The measures of the para-types are summarized in Table 1. Two of theparatypes have evident deformities: MCP 7957lacks all metatarsals and phalanges of the righthind foot; and MCP 7961 lacks the hallux of theright hind foot and has significant deformitiesof the head.

DISCUSSION

Although the phenetic groups in the genusMelanophryniscus are not well characterized, M.admirabilis is excluded from the M. tumifronsgroup because it lacks frontal swelling (Baldoand Basso, 2004). However, based on externalmorphological analysis, M. admirabilis appearsmore closely related to M. cambaraensis and M.macrogranulosus than to any remaining speciesin this genus. Thus, the morphology of the newspecies conflicts with the putative diagnosticcharacters of one of the phenetic species groupsof Melanophryniscus. A phylogenetic analysismust be conducted to test the reality of thesephenetic groups, posit the phylogenetic positionof the new species, and test the homology of thefrontal swelling.

Kwet et al. (2005) considered the state of RioGrande do Sul (Brazil) and the neighbor countryUruguay as the center of distribution of the genusMelanophryniscus. In fact, this is the ninth speciesreported in the state of Rio Grande do Sul (or10th, if M. devincenzii is considered), among thetotal of 19 known species in the genus. Six speciesoccur in Uruguay (Langone, 2003; Borteiro et al.,2005), of which three or four (depending on thestatus of M. devincenzii) also occur in Rio Grandedo Sul. Two other species recorded in Uruguayare probably new and currently are beingstudied.

M. admirabilis occurs along the southern slopesof the Brazilian Southern Plateau. All knownspecimens were caught during the day inside

FIG. 2. Dorsal view of the head of the holotype ofMelanophryniscus admirabilis sp. nov. (MCP 7923). Scalebar 5 5.0 mm.

FIG. 3. Ventral view of the right forelimb of theholotype of Melanophryniscus admirabilis sp. nov. (MCP7923). Scale bar 5 5.0 mm.

FIG. 4. Ventral view of the right hindlimb of theholotype of Melanophryniscus admirabilis sp. nov. (MCP7923). Scale bar 5 5.0 mm.

264 M. DI-BERNARDO ET AL.

marginal ponds or over flattened stone forma-tions along the left bank of the Forqueta River(Fig. 5), at 616 m above sea level. The vegetationis dominated by riparian native forest. Somenatural history observations indicate that thespecies may have reproductive activity duringOctober. In this month some clutches and tad-poles were recorded inside the marginal ponds,and many males were heard calling. Consideringthe apparent endemism of this species and thepaucity of data on its populational size, onlya small number of individuals were collected.About 75% of observed specimens were left inthe environment.

As in the case of other species in this genus(i.e., M. cambaraensis, M. macrogranulosus, Mela-nophryniscus orejasmirandai, M. tumifrons), M.admirabilis appears endemic to a very restrictedarea. Therefore, survival may depend on thedevelopment of a specific in situ conservationpolicy because of the high extinction risk thetaxa with small geographic ranges face (IUCNet al., 2004).

Acknowledgments.—We are in debt to thefollowing people that helped during fieldwork:

C. F. Batista, A. Escobar, C. Di-Bernardo, C. D. N.Castro, G. F. Pontes, R. B. Oliveira, R. L. Balestrin,L. H. Cappellari, J. Frota, C. Zank, W. Vaz, C.Schwingel, L. B. de Oliveira, and D. Goncalves.Raquel Rocha Santos made the illustrations. Weare grateful to the Curator of the AmphibianCollection of the Museu de Ciencias Naturais daFundacao Zoobotanica do Rio Grande do Sul,Marcio B. Martins, who allowed us to study thetype specimens of M. cambaraensis and M.macrogranulosus. Two anonymous reviewersmade improvements. We acknowledge GlauciaPontes and Roberto B. Oliveira for valuable helpduring laboratory work. MD is partially financedby the Conselho Nacional de DesenvolvimentoCientıfico e Tecnologico (Process CNPq #307.992/2004-7). RM has a doctoral fellowshipfrom Coordenacao de Aperfeicoamento de Pes-soal de Nıvel Superior (CAPES).

LITERATURE CITED

BALDO, D., AND N. G. BASSO. 2004. A new species ofMelanophryniscus Gallardo, 1961 (Anura: Bufoni-dae), with comments on the species of the genusreported for Misiones, Northeastern Argentina.Journal of Herpetology 38:393–403.

FIG. 5. Type locality of Melanophryniscus admirabilis sp. n. (Perau de Janeiro, Municipality of Arvorezinha,Rio Grande do Sul state, Brazil).

NEW BRAZILIAN MELANOPHRYNISCUS 265

BORTEIRO, C., F. KOLENC, M. TEDROS, AND F. GUTIERREZ.2005. Geographic distribution: Melanophryniscuspachyrhynus. Herpetological Review 36:199–200.

CESPEDEZ, J. A., AND M. MOTTE. 2001. Distribucion desapos del genero Melanophryniscus (Gallardo, 1961)en Argentina y Paraguay (Anura: Bufonidae).Boletın de la Asociacion Herpetolologica Espanola12:71–76.

CRUZ, C. A. G., AND U. CARAMASCHI. 2003. Taxonomicstatus of Melanophryniscus stelzneri dorsalis (Mert-ens, 1933) and Melanophryniscus stelzneri fulvogutta-tus (Mertens, 1937) (Amphibia, Anura, Bufonidae).Boletim do Museu Nacional 500:1–11.

DI-BERNARDO, M., R. B. OLIVEIRA, G. M. F. PONTES,J. MELCHIORS, M. SOLE, AND A. KWET. 2004. Anfıbiosanuros da regiao de extracao e processamentode carvao de Candiota, RS, Brasil. In E. C. Teixeiraand M. J. R. Pires (eds.), Estudos Ambientais emCandiota: Carvao e seus Impactos, pp. 163–175.FEPAM, Porto Alegre, Brazil.

ESCOBAR, A., R. MANEYRO, AND M. DI-BERNARDO. 2004.Rediscovery of Melanophryniscus macrogranulosus(Anura, Bufonidae), an endangered species ofAmphibia from the State of Rio Grande do Sul,Brazil. Biociencias 12:57–58.

FROST, D. R. 2004. Amphibian Species of the World: AnOnline Reference. Version 3.0 [cited 22 August,2004]. American Museum of Natural History, NewYork. Electronic database accessible at http://research.amnh.org/herpetology/amphibia/index.html.

GALLARDO, J. M. 1961. La ubicacion sistematica ydistribucion geografica de los Brachycephalidaeargentinos. 1a Reunion de Trabajos y Comunica-ciones de Ciencias Naturales y Geografıa del LitoralArgentino. pp. 205–212. Universidad Nacional delLitoral, Santa Fe, Argentina.

GRAYBEAL, A., AND D. C. CANNATELLA. 1995. A newtaxon of Bufonidae from Peru, with descriptionsof two new species and a review of phylogeneticstatus of supraspecific bufonids taxa. Herpetologica51:105–131.

HAAS, A. 2003. Phylogeny of frogs as inferred fromprimarily larval characters (Amphibia: Anura).Cladistics 19:23–89.

IUCN, CONSERVATION INTERNATIONAL, AND NATURESERVE.2004. Global Amphibian Assessment. http://www.globalamphibians.org. [Downloaded 15 Oc-tober 2004].

KWET, A., R. MANEYRO, A. ZILLIKENS, AND D. MEBS. 2005.Advertisement calls of Melanophryniscus dorsalis(Mertens, 1933) and M. montevidensis (Philippi,1902), two parapatric species from southern Braziland Uruguay, with comments on morphologicalvariation in the Melanophryniscus stelzneri group(Anura: Bufonidae). Salamandra 41:3–20.

LANGONE, J. A. 2003. Diversidad de la biota uruguaya.Anfibios. Anales del Museo Nacional de HistoriaNatural y Antropologıa 10:1–12.

LARSON, P. M., R. O. DE SA, AND D. ARRIETA. 2003.Chondrocranial, hyobranchial and internal oralmorphology in larvae of the basal bufonid genusMelanophryniscus (Amphibia: Anura). Acta Zoolog-ica 84:145–154.

Accepted: 20 March 2006.

APPENDIX

Examined Specimens

MCN: Museu de Ciencias Naturais da FundacaoZoobotanica do Rio Grande do Sul (Brazil). MCP:Museu de Ciencias e Tecnologia da Pontifıcia Uni-versidade Catolica do Rio Grande do Sul (Brazil).ZUFSM: Universidade Federal de Santa Marıa, RioGrande do Sul (Brazil). ZVCB: Facultad de Ciencias dela Universidad de la Republica (Uruguay).

Melanophryniscus atroluteus: ZUFSM 2474: Costa delrio Ibicuy, Manoel Viana, Rio Grande do Sul (Brazil).ZVCB 1678: Arrocera Conti, Artigas (Uruguay). ZVCB3137: Arrocera Conti, 25 km. W de Colonia Palma,Artigas (Uruguay). ZVCB 792, 6 km. SE de Melo, CerroLargo (Uruguay). ZVCB 10161: Camino de la lınea, 10km al NW de la Ciudad de Rivera, Rivera (Uruguay).

Melanophryniscus cambaraensis: MCN 9797 (holotype)and MCN 9760, MCN 9763-65, MCN 9768, MCN 9772,MCN 9776, MCN 9783, MCN 9786, MCN 9791, MCN9805, MCN 9810 (paratypes): Fortaleza dos Aparados,Cambara do Sul, Rio Grande do Sul (Brazil).

Melanophryniscus devincenzii: ZVCB 10661: ArroyoRubio Chico (Ruta 30), Rivera (Uruguay). ZVCB 10614:Est. Los Abuelos, Campos de FYMSA, Rivera, (Ur-uguay).

Melanophryniscus macrogranulosus: MCN 1694 (holo-type): Torres, Rio Grande do Sul (Brazil). MCN 1693,MCN 1695-99, MCN 1701-02 (paratypes): Torres, RioGrande do Sul (Brazil). MCP 7943: Barra do Ouro,Maquine, Rio Grande do Sul (Brazil).

Melanophryniscus montevidensis: ZVCB 7276-78: Al-vorada, Santa Vitoria do Palmar, Rio Grande do Sul(Brazil). ZVCB 4858-60: Balneario Perla de Rocha km258 Ruta 10, Rocha (Uruguay). ZVCB 10432-34: Perlade Rocha, Rocha (Uruguay).

Melanophryniscus orejasmirandai: ZVCB 7274-75: Sier-ra de Animas, Maldonado (Uruguay).

Melanophryniscus pachyrhynus: MCP 7953-55, MCP7962-63, MCP 7966-69: Fazenda Novosares, Sao Jer-onimo, Rio Grande do Sul (Brazil).

Melanophryniscus sanmartini: ZVCB 6376-77: Estable-cimiento Rocha, Rivera (Uruguay). ZVCB 8077: Esta-blecimiento Trinidad, Rivera (Uruguay).

266 M. DI-BERNARDO ET AL.