On the Identity of Species of Oreobates (Anura: Craugastoridae) from Central South America, with the Description of a New Species from Bolivia
The similar morphology of species in the genus Oreobates that inhabit rocky outcrops amidst the Seasonally Dry Tropical Forests and Cerrado savannas of central South America has led to taxonomic inaccuracies when only external morphology was considered. Therefore, researchers frequently fail to determine the true taxonomic status of some of these populations and thus attribute them to unidentified or misidentified species. Integrating different sources of data is of fundamental importance to understand the diversity of species in this group. Herein, we used external morphology (16 characters measured in 54 adult specimens), bioacoustics (238 advertisement calls of 19 specimens of Oreobates from six localities), and molecular evidence (based on 58 16S gene fragments representing 24 of the currently available names of Oreobates) to evaluate the taxonomic status and distribution range of Oreobates from 10 localities in central South America. We performed phylogenetic analysis, principal component analysis, and discriminant function analysis to evaluate the taxonomic status of Oreobates in central South America. We redescribe Oreobates heterodactylus based on a newly collected series of topotypes and additional material from other localities. We reassess the taxonomic status of Oreobates crepitans, herein considered as a junior synonym of O. heterodactylus. We also describe eastern Bolivian populations of Oreobates as a new species.ABSTRACT

(Left) Maximum likelihood phylogram based on 628 base pairs (bp) of mitochondrial 16SrRNA gene sequences from 58 individuals of Oreobates. Bayesian posterior probabilities and bootstrap values of >45 are shown on branches. Outgroup not shown for graphical purposes. (Right) Geographical distribution of Oreobates from 10 localities from eastern South America. Abbreviations: São Vicente, Santo Antônio do Leverger (SV; type locality of O. crepitans); Mata Fria, Chapada dos Guimarães (MF); Morro do Chapéu, Chapada dos Guimarães (MC); Estação Ecológica Serra das Araras, Porto Estrela (SA); Fazenda União, Rosário Oeste (RO); Dolina Água Milagrosa, Cáceres (DM); Gruta Fazendinha, Fazenda Hayashi, Cáceres (GF; type locality of O. heterodactylus); San Sebastián, Provincia Ñuflo de Chávez (SS); Provincia Chiquitos, Serranía de Santiago, Cerro del Arco (CA); Provincia Ángel Sandoval, Bella Boca (BB). Inset map: South America.

Variation in the relative proportions of tips of Fingers III and IV in populations of Oreobates from Mato Grosso, Brazil. (A) MNRJ 106A from Cáceres (GF; holotype of O. heterodactylus); (B) UFMT 19241 from Cáceres (GF); (C) UFMT 11119 from Porto Estrela (SA); (D) UFMT 19239 from Cáceres (GF); (E) MNRJ 3985 from Santo Antônio do Leverger, São Vicente (SV; paratype of O. crepitans). Finger expansions vary among specimens from different localities (e.g., B, C, and E) and among specimens from the same locality (e.g., A, B, and D).

Scatter plots of principal component analysis (PCA) component 1 versus PCA Component 2 scores of 16 morphometric (A) and six bioacoustic (B) variables for males of studied Oreobates populations from Brazil (Santo Antônio do Leverger, São Vicente [SV]; Chapada dos Guimarães, Mata Fria [MF]; Chapada dos Guimarães, Morro do Chapéu [MC]; Cáceres, Dolina Água Milagrosa [DM]; Cáceres, Fazenda Hayashi, Gruta Fazendinha [GF]; Porto Estrela, Estação Ecológica Serra das Araras [SA]; Rosário Oeste, Fazenda União [RO]) and Bolivia (Provincia Ñuflo de Chávez, San Sebastián [SS]; Provincia Ñuflo de Chávez, Cerro del Arco [CA]).

Oscillogram and spectrogram (DFT size = 512 samples) of the advertisement call of Oreobates from Brazil and Bolivia: (A) voucher call LH 62, from specimen UFMT 10058, recorded on 26 November 2009, 21:20 h, air temperature 24°C, municipality of Santo Antônio do Leverger, São Vicente (SV), state of Mato Grosso, Brazil; (B) voucher call LH 08, from specimen UFMT 11114, recorded on 18 December 2009, 00:20 h, air temperature 24.3°C, municipality of Porto Estrela, Estação Ecológica Serra das Araras (SA), Mato Grosso, Brazil; (C) voucher call LH 882, from specimen UFMT 19238, recorded on 14 November 2016, 19:40 h, air temperature 25.5°C, municipality of Cáceres, Fazenda Hayashi Gruta Fazendinha (GF), Mato Grosso, Brazil; (D) voucher call MJ-1683, from specimen SMF 88497, recorded on 25 November 2010, 21:16 h, air temperature 26.3°C, Provincia Ñuflo de Chávez, San Sebastián (SS), Department Santa Cruz, Bolivia. FD = dominant frequency.

Male topotype of Oreobates heterodactylus (UFMT 19241; SVL 25.0 mm) in dorsal, lateral, and ventral views.

Live specimens of Oreobates heterodactylus from Mato Grosso State, mid-western Brazil: (A) adult male (UFMT 19239) from the type locality, Gruta Fazendinha, Fazenda Hayashi, Cáceres (GF); (B) adult male (UFMT 19234) from Dolina Água Milagrosa, Cáceres (DM); (C) adult female (UFMT 19243) and (D) adult male (UFMT 19240) from Gruta Fazendinha, Fazenda Hayashi, Cáceres; (E) calling male (UFMT 19219) from São Vicente, Santo Antônio do Leverger (SV); (F) adult female (UFMT 11110) from Fazenda União, Rosário Oeste (RO).

Holotype of Oreobates chiquitanus sp. nov. (SMF 88497; male; SVL 24.6 mm): (A) dorsal and (B) ventral view; (C) lateral view; (D) ventral view of right hand, and (E) ventral view of right foot. Scale bar = 2 mm.

Live specimens of Oreobates chiquitanus sp. nov. from San Sebastián, Provincia Ñuflo de Chávez, Department Santa Cruz, Bolivia: (A) holotype (SMF 88497) and (B) adult female (SMF 88500).

Dorsal and ventral view of three paratypes of Oreobates chiquitanus sp. nov.: two adult males (SMF 88498-9) and one larger female (SMF 88500).
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